Health in Past Societies: Biocultural interpretations of human skeletal remains in archaeological contexts 9780860547167, 9781407348643

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Health in Past Societies Biocultural interpretations of human skeletal remains in archaeological contexts Edited by

Helen Bush and Marek Zvelebil

BAR International Series 567 1991

Published in 2019 by BAR Publishing, Oxford BAR International Series 567 Health in Past Societies © The editors and contributors severally and the Publisher 1991 The authors’ moral rights under the 1988 UK Copyright, Designs and Patents Act are hereby expressly asserted. All rights reserved. No part of this work may be copied, reproduced, stored, sold, distributed, scanned, saved in any form of digital format or transmitted in any form digitally, without the written permission of the Publisher. ISBN 9780860547167 paperback ISBN 9781407348643 e-book DOI https://doi.org/10.30861/9780860547167 A catalogue record for this book is available from the British Library This book is available at www.barpublishing.com BAR Publishing is the trading name of British Archaeological Reports (Oxford) Ltd. British Archaeological Reports was first incorporated in 1974 to publish the BAR Series, International and British. In 1992 Hadrian Books Ltd became part of the BAR group. This volume was originally published by Tempvs Reparatvm in conjunction with British Archaeological Reports (Oxford) Ltd / Hadrian Books Ltd, the Series principal publisher, in 1991. This present volume is published by BAR Publishing, 2019.

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CONTENTS

L i s to fC on t r ibu to r s L i s to fF igu res L i s to fT ab les A cknow ledgemen t s P ref ace

I V v i v i v i i

PART 1 : Concep ts a nd Me thods 1 .

P a tho logy a nd h ea l th i np a s ts oc ie t ie s : a ni n t roduc t ion . H e len B ush a nd Marek Z ve leb i l

3

2 .

C oncep t so fh ea l th a nd s t re s s . H e len B ush

1

3 .

A ccoun t ing f o r t h e e vo lu t ion o fp sychosoma t ic p h enomena : d id u nco o pe ra t ive U pp e rP a la eo l i th ic p eop le b ecome i la nd d i e? M ichae lE .H y land a nd W in S cu t t

2 3

4 .

H ea l th , a dap t a t ion a nd ma ladap t ion i np a s ts oc ie t ies . A lan G oodman

3 1

5 .

D ir ec t g ene t ic a nd i mmuno log ica l i n fo rma t ion i nt h e r e con s truc t ion o f h ea l th a nd b i ocu l tu ra l c ond i t ion s o f p a s t p opu la t ion s : an ew p ro spec t f o ra r chaeo logy . C r i s t ina C a t taneo

3 9

6 .

D en ta l h i s to logy a sa ni n dex o fp a s tc ommun i ty h ea l th . S imon H in son

5 3

PART 2 : Case S tud ies 7 .

P a t te rn so fl i f e a nd d ea th : t h ep a laeodemog r aphy o f Med ieva lY o rk . A nne G rauer

6 7

8 .

E p idem io log ica l s t ud ie s o f d en ta l e name l h ypop la s ia s i n Mex ico a nd B radford : t h e ir r e levan ce t oa r chaeo log ica ls ke le ta ls t ud ie s . K e i th Dob l zey a nd A lan G oodman

8 1

9 .

A naem ia i nR oman B r i ta in : P oundbury C amp . P a t ty S tuar t -Macadam

1 0 1

1 0 .

I n te rp re t ing m ea su re s o fc ommun i ty h ea l th d u r ing t h e L a te P r eh is to r ic p e r iod i nM idd le T enne s see : ab iocu l tu ra l a pp roach . L es l ie E i senberg

15

1 .

H ea l th s t a tu s o fE u rop ean p opu la t ion s a tt h e a g r icu l tu ra l t r an s i t ion a nd t h e i mp l ica t ion s f o rt h e a dop t ion o ff a rm ing . C hr i s topher Me ik l e john a nd Marek Z ve leb i l

1 29

1 1

L IST OF CONTRIBUTORS

H e len B u sh ,P h .D . D epa r tmen to fA r chaeo logy a nd P r eh i s to ry U n ive r s i ty o fS hef f ie ld . C r i s t ina C a t taneo ,M .A . D epar tmen to fA r chaeo logy a nd P r eh i s to ry U n ive r s i ty o fS hef f ie ld . K e i th D obney , M .Sc . D epa r tmen to fA r chaeo log ica l S c ience s U n ive r s i ty o fB rad fo rd . L e s l ie E .E isenbe rg ,P h .D . D epa r tmen to fA n th ropo logy N ew Y o rk U n ive r s i ty . A lan G oodman ,P h .D . S choo lo fN a tu ra l S c ience H amp sh i re C o l lege Amhe r s t , Ma ssa chu se t t s . A nne G raue r ,P h .D . D epa r tmen to fA n th ropo logy L oyo la U n ive r s i ty o fC h icago . S imon H i l lson ,P h .D . I n s t i tu te o fA r chaeo logy U n ive r s i ty C o l lege L ondon . M ichae lH y land ,P h .D . D epa r tmen to fP sycho logy P lymou th P o ly techn ic . C h r is tophe r Me ik le john ,P h .D . D epa r tmen to fA n th ropo logy U n ive r s i ty o fW inn ip eg C anada . W in S cu t t ,B .A . P lymou th C i ty Museums P lymou th . P a t ty S tua r t - Macadam ,P h .D . D epa r tmen to fA n th ropo logy U n ive r s i ty o fT o ron to . Ma rek Z ve leb i l ,P h .D . D epa r tmen to fA r chaeo logy a nd P r eh i s to ry U n ive r s i ty o fS hef f ie ld .

i v

L IST OF F IGURES 2 . 1 4 . 1

4 . 2 5 . 1

5 . 2 5 . 3 5 . 4 5 . 5

6 . 1

6 . 2 6 . 3 6 . 4

6 . 5 6 . 6

7 . 1 7 . 2 7 . 3 8 . 1 8 . 2 8 . 3 8 . 4 8 . 5 8 . 6 8 . 7a 8 . 7b 8 . 8a 8 . 8b 8 . 9 8 . 10

9 . 1 9 . 2

T he G ene ra lA dap ta t ion S ynd rome ( f rom S e lye , 1957 ) . Ag ene ra l mode l f o rs t udy ing t h er e la t ion sh ip b e tween h ea l th , s t r e s s , a nd a dap ta t ion i n s k e le ta l p opu la t ion s ( f rom G oodman a nd A rme lago s , 1 989 ) . Ap ea san tf am i ly f r om T ezon t eopan , Mex i co . S cheme s how ing t h e i n t roduc t ion o fg ene t ic ( a long w i th e nv i ronmen ta l ) i n fo rma t ion i n t h e r e con s t ruc t ion o f t h e h ea l th s t a tu s o f as k e le ta l p opu la t io n . D raw ing o f ac l a s s I HLA a n t igen ( a f te rA lbe r t se ta l , 1 983 ) . S chema t i c e xamp le o f t wo h ap lo type s i na n i n d iv idua l ( a f te r S ch ieb l , 1 986 ) . I n he r i tan ce u n i t si n af am i ly s e g r ega t ion a na ly s i s( a f t e rB ende r , 1 984 ) . G ene r a l s c heme s how ing t h e u s e o f g ene t i ca l ly i n fo rma t ive mo lecu le s , e x t ra c ted f r om s k e le ta l r ema in s , i nt h e r e con s t ruc t ion o f t h e d ynam ic s a nd c u l tu r eo f ap opu la t ion . I n c remen ta l s t ruc tu r e s i nt h e d en t ine o f ah uman c an ine f r om t h e s i te o f K e rma , N ub ia ( c . 1 720 1550 B .C . ) . Ma te r ia l f r om t h e D uckwo r th L abo r a to ry ,U n ive r s i ty o fC amb r idge . R ange s o f d en t ine f o rma t ion t i me s f o r v a r iou s h uman p e rmanen t t e e th ( d a ta f r om G u s ta f son a nd K o ch , 1 974 ) . P r i sm s i nt h e e name l o f ah uma n f i r s t m o la r f r om t h e C oppe rga te s i te , Y o rk . B rown s t r iae o fR e tz iu s i nt h ee name lo f ah uman c an ine f r om t h e s i t eo f K e rma , N ub ia ( c . 1 750 -1550 B .C . ) . Ma te r ia l f r om t h e D uckwo r th L abo r a to ry ,U n ive r s i ty o fC amb r idge . R ange s o f e name l f o rma t ion t i me s f o r v a r iou s h uman p e rmanen t t e e th ( d a ta f r om G u s ta f son a nd K o ch , 1 974 ) . P e r ikyma ta o n t h e c r own s u r f a ce o f ah uman c an ine f r om t h e L ive rpoo l S t ree t /B r oad S t r ee t s i te , L ondon . Ma te r ia l f r om t h e E nv i ronmen ta l A r chaeo logy U n i t , Mu seum o fL ondon . Mo r ta l i ty r a te f o rt h eS t . H e len -on the - Wa l l sp opu la t ion . Mo r ta l i ty r a te b ys e xi nt h eS t .H e len -on the - Wa l l sp opu la t ion . S u rv ivo r sh ip c u rve b y s e xi nt h eS t .H e len -on the - Wa l l sp opu la t ion . F requen cy o f DDE s f r om c h i ld r en e xam ined a t t e n s c hoo l s i n t h e B rad fo rd r e g ion . F requen cy o fL EH f r om c h i ld r en e xam ined a tt e n s c hoo l si nt h e B rad fo rd r e g ion . B rad fo rd L EH ,m ax i l la ry t e e th F requen cy o f DDE s i nE u ropean a nd A s ian c h i ld ren f r om U nde r c l i f f e a nd G t . H o r ton m idd le s c hoo l s . F requen cy o fL EH a nd p i t s i nA s ian c h i ld r en f r om U nde r c l if f e a nd G t . H o r ton m idd le s c hoo l s . F requen cy o f d i f f e r en t t y pe s o fe name l d e f ec t s i nE u ropean a nd A s ian c h i ld r en f r om U nde r c l i f f ea nd G t . H o r ton m idd le s c hoo l s . T ezon teopan -p i t h ypop la s ia o f m ax i l la ry t e e th , s upp lemen ted v s . n on s upp lemen ted . T ezon teopan -p i th ypop la s ia o fm and ibu la rt e e th , s upp lemen ted v s . n on s upp lemen ted . T ezon teopan - L EH o f m a x i l la ry t e e th , s upp lemen t ed v s . n on s upp lemen ted . T ezon teopan - L EH o f m and ibu la r t e e th , s upp lemen ted v s . n on s upp lemen ted . T ezon teopan -L EH o f m a x i l la ry f i r s t i n c i so r s t e e th , s upp lemen ted v s . n on supp lemen ted . H ypo the t ica l m ode l s how ing d i f f e r en t p a t te r n s o f L EH p roduced b y i d en t ica l s t r e s so r e ven t s i n i n d iv idua l s /popu la t ions o f v a ry ing h ea l th s t a tu s . V au l tl e s ion so fp o ro t ic h ype ro s to s i s . O rb i ta ll e s ion s( c r ib r ao rb i t a l ia )o fp o ro t ic h ype ro s to s i s( P e ruv ian o r ig in ) .

2 1 3 7

3 8 5 0

5 0 5 1 5 1 5 2

5 9

6 0 6 1 6 2

6 3 6 4

7 8 7 9 8 0 9 2 9 2 9 3 9 4 9 4 9 4 9 5 9 6 9 7 9 8 9 9 1 00

1 08 1 08

9 . 3a 9 . 3h 9 . 4 9 . 5 9 . 6 9 . 7 9 . 8 9 . 9 9 . 10 9 . 11 1 0 .1 1 0 .2 1 0 .3 1 0 .4 1 0 .5 1 0 .6 1 0 .7 1 0 .8 1 0 .9 1 0 .10 1 0 .11 1 .1

I lu s t ra t ions o fl e s ion t y p e s . D e sc r ip t ion o fl e s ion sa nd a r ea so fo r b i t . Map o fD o r che s t e rs how ing P oundbu ry C amp l o ca t ion . G rave d i s t r ibu t ion a tP oundbu ry C amp . A ge d i s t r ibu t ion f o rP oundbu ry C amp i n d iv idua l s . D i s t r ibu t ion o fc r ib ra o rb i ta l ia w i th a ge . D i s t r ibu t ion o fv au l tl e s ion sw i th a ge . O rb i ta ll e s ion t yp ei nP oundbu ry C amp i n d iv idua l s . V au l tl e s ion t y p ei nP oundbu ry c amp i n d iv idua l s . L oca t ion a nd t y p eo fc r ib ra o rb i ta l ia . Map o ft h eS t a te o fT enne s se e . P lan v i ew o ft h eA ve rbuch s i te , i n c lud ing c eme t e r ie s ,s t ru c tu re s , f e a tu r e s a nd d i s tu rbance f r om mode rn c on s t ruc t ion a c t iv i t ie s( Reed , 1 984 :8 ) . At y p i ca lM idd le C umbe r land s t one b ox g r ave . H ea led p e r io s to s i s a l ong t h e e n t i r e s h a f t o f t h e t i b ia o f B u r ia l 5 17B , a f ema le ,a ged 3 0 -34 y ea r s ,f r om t h eA ve rbuch s i t e . R e la t ive f r equen c ie so fa c t ive a nd h ea led p ro l i f e ra t ive t i b ia ll e s ion si nt h e A ve rbuch p opu la t ion . R e so rp t ive l e s ion s a nd c o l lap se o f t h e t h o ra c ic s p ine o f a n A ve rbuch f ema le ,a ged 3 0 -34 y ea r s . L a rge , a c t ive p o ro t ic l e s ion s o n t h e p o s te r io r o f t h e c r a n ia l v au l t i na n A ve rbuch f ema le ,a ged 1 8 .5 -19 .5 . R e la t ive p e r cen tage s o f a c t ive a nd h ea led p o ro t ic l e s ion s a c ro s s a l l A ve rbuch a ge g roup s . B i la te r ia l o rb i ta l l e s ion s , w h i ch e xh ib i t ac omb ina t ion o f h ea led a nd a c t ive s t a te s ,i n ay oung c h i ld ( 3 .5 -4 .5 y ea r s )f r om A ve rbu ch . R e la t ive p e r cen tage s o f a c t ive a nd h ea led o r b i ta l l e s ion s a c ro s s a l l A ve rbuch a ge g roup s . Mo r ta l i ty p ro f i le o ft h eA ve rbuch p opu la t ion ( Be r ryman , 1 981 :60 ) . C eme te r ie sa nd B u r ia l si n Me so l i th ic E u rope .

1 09 1 09 10 10 11 11 12 12 13 13 1 2 1 1 22 1 23 1 24 1 24 1 25 1 25 1 26 1 26 1 27 1 27 1 4 5

L IST OF TABLES 2 . 1 5 . 1 5 . 2a 5 . 2b 5 . 3 7 . 1 7 . 2 7 . 3 7 . 4 7 . 5 8 . 1 8 . 2 9 . 1 1 .1

T he a n th ropo log ica lu s age o f' s t r e s s ' . L i s to f HLA a n t igen s( a f t e rB ende r , 1 984 ) . HLA -A ,B , Ca l le le f r equenc ie s( % )( a f te rB end e r , 1 9 84 ) . S ome HLA -A a nd B a n t igen f r equenc ie s ( % ) a mong E u ropean s , M idd le E a s te rn s ,I n d ian s , Mongo lo id s ( a f te rD au s se ta nd C o lomb an i , 1 973 ) . S ome s i gn i f ican t HLA a n t igen a nd d i sea se a s so c ia t ion s . L if ef o rt h et o ta lp opu la t ion . A s soc ia t ion b e tween b u r ia la r ea a nd a ge a td ea th f o rs ubadu l t s . L if et a b le sb yg ende r . K o lmogo rov -Sm ir nov t e s t f o r d i f f e r ence s b e tween a ge a t d ea th r a te s b y g ende r . P e r cen tage o fa du l tm a le sa nd f ema le si nt h et o ta lp opu la t ion . S e le c ted p r ev iou ss t ud ie so f DDE s . F requency o fd en ta le name ld e f ec t si nB rad fo rd b ys c hoo la nd w a rd . I n c idence o fc r ib ra o r b i ta l ia a nd v au l tl e s ion sw i th a ge . C a r ie sr a t e si nt h eE u rop ean Me so l i th i c .

2 0 4 6 4 7 4 8 4 9 7 5 7 6 7 6 7 7 7 7 9 0 9 1 1 0 7 1 4 4

ACKNO WLEDGE MENTS We t h ank R u s se l lA dam s ,o fJ . R .C o l l i sP ub l ica t ion s ,f o rp r epa ra t ion o ft h ec ame ra r eady c opy o ft h i s v o lume . We a l so t h ank Mav i s T o r ry a nd E dd ie Mo th f o r t h e i r h e lp i nt h e p r epa ra t ion o f t h e t e x t a nd i l u s t ra t ion s . We a r eg r a te fu lt oo u rc on t r ibu to r sf o rt o le r a t ing t h ep ro t r a c ted c omp le t ion t i me o ft h i sv o lume ,w h ich a r o se f r om c i r cums tance sb eyond o u rc on t ro l .

v i

PREFACE T he t h eme o f t h i s v o lume i sh ea l th : i t s r e cogn i t ion i n , a nd s i gn i f ican ce f o r , p a s t s o c ie t ie s . T he v o lume h a s i t s o r ig in s i nt h e s e s s ion ' A Ma t t e ro fL i f e a nd D ea th ' w h ich t h ea u tho r s o rgan i sed a sp a r to f t h e T heo re t ica l A r chaeo logy G roup ( T .A .G . ) C on f e r ence a tt h e U n ive r s i ty o f S he f f ie ld i n D e cembe r 1 988 . T he p ape r sp r e sen ted a tt h a ts e s s ion a r ea ugmen ted h e r eb yf o u ra dd i t iona lc on t r ibu t ion s . T he v o lume i sd i v ided i n to t wo h a lve s ,t h ef i r s tp r e sen t ing c oncep t so fh ea l th a nd i n t roduc ing s ome o f t h e w ay s i nw h ich t h e c ond i t ion c an b e i d en t i f ied i np a s tp opu la t ion s . T he s e cond h a lf c on s i s t s o ff i ve c a se s t ud ie s ,p r inc ipa l ly o fE u ropean m a t e r ia l . O u r i n t roduc t ion e xam ine s t r ad i t iona l a pp roa che s t o t h e s t udy o f h uman s ke le ta l r ema ins f r om a r chaeo log ica l c on tex t s , a nd c ompa re s t h em t ot h e ' b io cu l tu r a l ' a pp roa ch w h ich h a s d eve loped i nN o r th Ame r ica . T he ma in f e a tu re s o ft h i s c ompa ra t ive ly n ove l a pp roa ch a r e p r e sen ted a nd t h e s i gn i f icance o f h ea l th a tp opu la t ion l e ve l i sc on s ide red . F o l low ing t h e i n t roduc t ion , t h ec hap t e rb y H e len B u sh l o ok s a t d e f in i t ion so fh ea l th ,a nd i np a r t i cu la rt h ee co log ica ld ef in i t ion u pon w h ich t h eb i o cu l tu ra la pp roa ch r e s t s . T he c hap te rt r a ce s t h e p r og re s s ion f r om d e f in i t ion so fh ea l th t ot h e i n te rp r e ta t ion o ft h e c ond i t ion i np a s t p opu la t ion s t h rough t h e m ed ium o f ' i nd ica to r s o f s t r e s s ' . U s ing c l in ica l s t ud ie s a s e xamp le s , B u sh d emon s t ra t e s t h a t p sycho log ica l f a c to r s a r ec apab le o fa f f e c t ing g r ow th a nd p ropo se s t h a t t h ey m ay t h u s i nf luence t h e f o rma t ion o f ' s t r e s s ' m a rke r s , s i n ce t h e se r ef lec t g r ow th d i s tu rbance . T he l i nk b e tween m o rb id i ty a nd p sycho log ica ls t a te si sa l so m ade b yM i cha e lH y land a nd W in S cu t t ; i nC hap te rT h ree t h ey p r e sen t af u nc t iona lt h eo ry o fi l ne s st oa c coun tf o rt h er e la t ion sh ip b e tween t h e se t wo f a c to r s . A lan G oodman ' s c hap te rc on t inue s t h e e xp lo ra t ion o ft h e n o t ion o fh ea l th a nd o fi t s s i gn i f icance f o r p a s ts oc ie t ie s . H ed emons t ra te s t h ec onne c t ion b e tween h ea l th a nd a dap ta t ion t h rough a n e xam ina t ion o f t h ep henomenon o fs ma l lb ody s i ze . I nC hap te r F ive , C r i s t ina C a t taneo e xp la in s h ow g ene t i c f a c to r s i nf luence d i sea se s u s cep t ib i l i ty , a nd d e sc r ibe s r e sea r ch a imed a tr e t r iev ing m o le cu la r i n fo rma t ion f r om s k e le ta l m a t e r ia l i n o rde r t h a t t h e g ene t ic s u s cep t ib i l i ty t od i sea se o f ap opu la t ion c an b ec on s id e r ed w hen i t sa e t io logy u nde ri n ve s t iga t ion . P a r tO ne i sc omp le t ed b y S imon H i l l son ' s c on t r ibu t ion , w h ich d e sc r ibe s t h e u t i l i ty o fh i s to log i ca l s t ud ie s o fa nc ien tt e e th . T he p u rpo se o f t h e f i ve p ape r s i n P a r t Two i st o d emon s t r a t e t h e b i o log ica l a pp roa ch a nd t h e a pp l ica t ion o fc oncep t s w h i ch w e r e i n t roduced i nP a r t O ne . I nC hap t e r S ev en , A nne G r aue r f o cu se s o n t h e c on t rove r s ia l s ub jec to fp a laeodemog r aphy . H e r s t udy o f al a rge s k e le ta l a s semb lage f r om m ed ieva l Y o rk c omb ine s a r chaeo log ica l , h i s to r i ca l a nd p a laeodemog r aph i c d a ta i no rde r t oi l u s t r a te t h e p i t f a l l s a nd p o ten t ia la dvan tage so ft h i sf i e ld o fs t udy . T he n ex t t wo c hap t e r s e a ch c oncen t ra te o n as i ng le s k e le ta l i n d i ca to r o f h ea l th s t a tu s . H i l l son i n t roduced t h e d en ta l l e s ion s k nown a s e name l h ypop la s ia i nC hap te r S i x , a nd i nC hap te r E igh t K e i th D obney a nd A lan G oodman r e po r t o n e t hnog raph ic s t ud ie s o f t h i s l e s ion f r om t wo v e ry d i f f e ren t l o ca t ions , B rad fo rd a nd Mex i co . C omp lex i t ie s a r e r e vea led w h ich g i ve t h e l i e t o o ve r s imp l i f ied i n te rp re ta t ion s o f e name l h ypop la s ia i n a r chaeo log ica l a nd s k e le ta l a s semb lage s . I n t h e f o l low ing c hap t e r , P a t ty S t ua r t -M acadam p r e sen t s t h e r e su l t s o f h e r s t udy o f p o ro t ic h ype ro s to s i s ( a c ond i t ion i n d ica t ive o f a naem ia ) i n t h e s k e le ta l a s semb lage f r om P oundbu ry C amp , aR omano -B r i t i sh s i te i n D o r se t . T he c hap te ri n c lude s ad i s cu s s ion o ft h ec omp lex a e t io logy o ft h ec ond i t ion . I nC hap te rT en ,L e s l ie E i senbe rg d e s c r ibe sh e rb i ocu l tu ra la na ly s i so ft h ed i sea se w h ich s he n o ted i na s k e le ta l a s semb lage f r om A ve rbu ch , ap r eh i s to r ic v i l lage s i te i nw ha ti sn ow T enne s see . H e rc hap te ri sa c l ea re xpo s i t ion o ft h ew ay i nw h ich h ea l th s t a tu sc an b eu s ed t op o s tu la te c au se so fc u l tu re c hange . T he f i na l c hap te r i nt h e v o lume i sb y C h r i s tophe r Me ik le john a nd Ma r ek Z ve leb i l , w ho s e t o u t t o s how h ow ab i o cu l tu ra l a na ly s i s o f s k e le ta l r ema in s c an s h ed l i gh t o n t h e t r an s i t ion f r om h un t ing a nd g a the r ing t of o od p roduc t ion i nE u rope . T he i n t roduc to ry c hap te re xp la in s o u r m o t iva t ion i nb r ing ing t o ge the r t h e se c on t r ibu t ion s . T he f i na l s e n tence o ft h e v o lume e xp r e s se s o u r h ope s f o r i t : ' I f( t he v o lume ) h a s s u cce ed ed i nd emons t ra t ing t h e p o t en t ia la nd t h ev a lue o ft h eb i o cu l tu r a la pp roa ch ,t h en i tw i l lh ave s e r ved i t sp u rpo se ' . H e len B ush a nd Marek Z ve leb i l S hef f ie ld F ebruary , 1 991 .

v i i

PART 1

C oncep t s a nd Me thods

1 .

PATHOLOGY AND HEALTH I N PAST SOC IET IES : AN I NTRODUCT ION .

H e len B ush a nd Marek Z ve leb i l l a ck c l in ica l t r a in ing t o v en tu r e i n to t h e i n f in i t e ly s ub t le f i e ld o f a nc ien t d i sea se " ( We l l s , 1 964 :20 ) .

Approaches t o P a laeopa tho logy a nd Hea lth i n B r i t ish a nd Amer ican P rac t ice

" A l though b e ing a d o c to r i s a n ece s sa ry c ond i t ion , t h e c ond i t ion i sn o t s u f f ic ien t . I ti sn e ce s sa ry , a sw e l l , t og e t a g ood a n th ropo log ica l f o unda t ion i n c lud ing , i f p o s s ib le , a g ood a r chaeo log ica l b a ckg round , n o t o n ly t o e s cape t h e p seudopa tho logy p i t f a l l s b u t , o ve ra l l , t or e a l ize t h e n ece s sa ry c o l labo ra t i on w i th a r chaeo log i s t s " ( Da s tugue , 1 980 :3 ) .

T he s ke le ta l r ema in s o fp a s tp opu la t ion s h ave b een a f o cu s o f i n te r e s t f o r p hy s ica l a n th ropo lo g i s t s a nd m ed ica l p r a c t i t ione r s s i nce t h e n i ne t e en th c en tu ry . P a ra l le l l i ne s o f i n ve s t iga t ion e x i s ted a tt h i s t i me : p hy s i ca l a n th ropo log i s t sw e r e c h ief ly c once rned w i th m e t r ic a na ly s i s ( Wa shbu rn , 1 953 ) , w h i le t h e a t ten t ion o f o t he r s c ien t i s t s w a s t u rned t op a tho log ica l c ond i t ion s . T he t e rm " p a laeopa tho logy " w a s c o ined i n d ependen t ly i n Ame r i ca a nd E u rope , a nd i tw a s S i r Ma r c A rmand R u f f e ri n1 9 13 w ho , d e s c r ib ing h i s s t udy o f d i sea se i n E gyp t ian m umm ie s , d e f ined p a la eopa tho logy a s " t he s c ience o f t h e d i sea ses w h ich c an b ed emons t r a ted i nh uman a nd a n ima l r ema in s o f a nc ien t t i me s " ( Mood ie , 1 923 :21) . T he w o rk o fR u f f e r a nd o fo t he r s , s u ch a s Mood ie a nd E l l io t -Sm i th , w a s e s sen t ia l ly c l a s s if ica to ry ( O r tn e r , 1 988 ) , t h e b roade r i mp l i ca t ion s o f d i sea se g o ing u n con s ide r ed . T he e l uc ida t ion o f t h e n a tu ra l h i s to ry o f s p ec i f ic d i sea se sc on t inue st ob et h eg oa lo fm any s t uden t s o fs ke le ta lr ema in s :

T he i n f e r ence i st h a t " t he n ece s sa ry c o l labo ra t i on w i th a r chaeo log i s t s " i s ao ne -way p roce s s t o t h e b ene f i t o f t h e p a laeopa tho log i s t . I ti ss ma l l w onde r t h a t m any a r cha eo log i s t s a r e i n d if f e ren t , d i sm i s s ive a nd s ome t ime s e ven h o s t i le t owa rd s h uman s k e le ton sa nd t h e i re xcava t ion . Af u r the r e xp lana t ion f o r s u ch u nf avou rab le a t t i tude s m ay b e t h a t o fi g no rance . O ne o f u s r e c en t ly s p en t ap e r iod o ft i me e xcava t ing a n u r ban c eme te ry , w h ich p r e sen ted a n e xce l len t o ppo r tun i ty t oe l i c i tt h eo p in ion so ft h o se a c t ive ly e ngaged i n e xcava t ion . O ne s u ch , w ho se u ncove r ing o f a n a do le s cen t s ke le ton w a s m e t icu lou s , a dm i t t ed t oh av ing n o i d ea w ha t i t s s ub sequen t s t udy c ou ld r e vea l . O the r s r e f e r red i n ev i tab ly t ot h e d e t ec t ion o f d i sea se , a nd t ot h e a s se s smen to fa ge -a t -dea th a nd g ende r . U nawa r e o f t h e p o ten t ia l o f h uman s ke le ta l r ema in s , m any a r chaeo log i s t s v i ew t h em a s , a t b e s t ,a n i re levance ,a nd w hen e n coun te r ed i ns i tu a s o b je c t s w ho se e xcava t ion i st i me -consum ing a nd w h ich s omehow d oe s n o t c on s t i tu te " r ea l " a r chaeo logy . I mp l i c i t ly o r e xp l ic i t ly , s ke le ta l r ema in s a r e h e ld t ob e i n f e r io r t oo t he r a r tef a c t s , e ven t h ough t h e f o l ly o fp ay ing m o r e a t ten t ion t o t h e f i bu la w h ich f a s ten s t h e s h roud t h an t o t h e c o rp se i t s e l f h a s b een e xpo sed ( Denne l l , 1 979 ) . T he re a r e , o f c ou r se , n ume rou s B r i t i sh a r chaeo l o g i s t s w i th e x t en s ive e xpe r ience o f s ke le ta l e xcava t ion a nd i n fo rmed e n thu s ia sm f o r t h e i r a na ly s i s . E ven s o , w e w e re a mu sed b y o ne w ho u ncon sc iou s ly b e t rayed h imse l f w hen r e coun t ing t h e t h e f t o f s ku l l s f r om t h e s i te w h ich h e w a s d i re c t ing : f o r tuna te ly , h e r ema rked , n one o f t h e a r t ef a c t sw a ss t o len ! T he l i m i ted c oncep t o f h uman s ke le tons m ay b e a t t r ibu ted t ot h e w ay i nw h ich t h ey h ave b een r e po r ted i nt h e p a s t , a s a ppend ice s t oe xcava t ion r e po r t s , r a the r t h an a s a n i n teg ra l p a r t o f t h e

" . .ske le ton s c an y i e ld u se fu l i n fo rma t i on o n t h e p r e sence o r a b sence a nd i n c idence o f c e r ta in b one a nd j o in t d i sea se s i n o u r a nce s to r s a nd c an a dd v a luab le h i s to r ica l p e r spe c t ive t o o u r s t ud ie s o f t h e v a r iou s d i sea se s a ff l ic t ing m ank ind t o day " ( Rog e r s a nd D ieppe , 1 983 :1194 ) . We w ou ld a r gue t h a t t h is c l in ica l e mpha s i s g oe s s ome w ay t oe xp la in ing t h e l a ck o f i n te r e s t o fm any B r i t i sh a r cha eo log i s t s i nh uman s k e le ta l r ema ins ; a r chaeo log i s t s h ave b een a c t iv e ly d i s c ou r aged f r om i n vo lvemen ti nt h e i rs t udy : " The r e h a s b een a l o t o f c on fu s ion b ecause m any o f t h o se s t udy ing s k e le ta l m a te r ia l h ave b een a r chaeo log i s t s o r a n th ropo log i s t s w i th n o m ed ica l b a ck g round s o t h a t s i gn i f i can t b ony c hange m ay b e m i s sed o rt e x tbook d e s c r ip t ion s o f d i sea se m i sapp l ied ( Roge r s a nd D ieppe , 1 983 :1192 ) . " . .a p ro found c l in i ca l k now ledge i s n eeded t os o lve t h e p rob lems t h a ta r e m e t . I ti s mo s t u nw i se f o r a n th ropo log i s t s w ho

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d i vo r ced f r om e a ch o t he rc oncep tua l ly ( s ee B u sh , t h i s v o lume ) . I ti sr a the r t h a t t h e a r ch aeo log i s t , c once rned w i th t h e " i l lum ina t ion o f h uman s o c ie ty a nd i t s d ynam ics " ( A cke rne ch t , 1 9 53 :126 ) , s e ek s ab r oade r v i ew t h an af o cu s o n d i sea se p e r s ec an p rov ide . A cco rd ing t oO r tne r ( 1 988 ) t h i s i sn ow b e ing r e cogn i sed i n c r ea s ing ly b yp a la eopa t ho log i s t s w ho h ave b egun t oi n qu i re i n to t h e s i gn i f ican ce o f t h e d i sea se s w h ich t h ey d e te c t s k e le ta l ly , r a the r t h an s imp ly a temp t ing t h e i r d i agno s i s . S o l o ng a st h e l a t t e rr ema in s t h e g oa l , t h e a r cha eo log i ca l p o ten t ia l o f h uman s k e le ta lr ema in si sw a s t ed . I n ev i tab ly , t h e r e a r e d i f f i cu l t i e s a nd d i sag r eemen t sw hen s k e le ta lr ema in sa r ea na ly sed i nt h i s w ay . T he w ide ra pp l ica t ion a nd e va lua t ion o fi n d ica to r s o f s t r e s s h a s r e vea led t h a t i ns ome c a se s t h e i ri n te rp r e ta t ion i nt e rm s o ft h e h e a l th o f p a s t p opu la t ion s h a s b een n a ive . A c l a s s ic e xamp le i st h a t o ft h e t r an sve r se l i ne s w h i ch a r e f r equen t ly o b se rved o n t h e r a d iog r aph s o f l o ng b one s , a nd w h i ch a r e c ommon ly r e f e r r ed t o a s H a r r i s l i ne s . T he se w e re h e ra lded a s " a n ew a pp roa ch t oa n c ien t d i sea se " (We l l s , 1 9 61 :526 ) . W i th h i s c u s toma ry e l oquence , We l l s ( 1 967 :390 ) s ummed u p t h e p i c tu r e w h ich t r ad i t iona lp hy s i ca l a n th ropo logy p a in t sf o ru s : " . .we c an s e ea ne a r ly i n d iv idua l a to n ly o ne m omen t -t h e m omen t o f d ea th . I ti s as n ap sho t , n o t am ov ing p i c tu r e " . I t w a si no rde rt om ove f r om t h i s s t a t i cp o s i t ion t h a t h e p romo ted t h e r a d io log i ca l e xam ina t ion o f a r chaeo log i ca l s k e le ton s f o r t h e p r e s ence o f H a r r i s l i ne s ; t h ey w ou ld b e as t ep t owa rd s a d i a ch ron i c p o r t r aya l o f t h e l i f e o f a n i n d iv idua l , b e cau se i tw a s b e l ieved t h a t t h ey r ef l e c t h ea l th s t a tu s a t d i f f e r en t s t age s d u r ing t h e y e a r s o f g r ow th , a s o ppo sed t oh ea l th a t t h e m omen t o f d ea th ,i fs u ch as t a t ec an b es a id t oe x i s t ! We l l s u s ed H a r r i s l i ne s t o m ak e s o c ia l i n f e r ence s . F o r e xamp le , f r om a n i n te r -popu la t i ona l s t udy i n w h ich t h e r e la t ion sh ip b e tween g ende r , a ge -a t -d ea th a nd H a r r is l i ne f r equen cy w a sa s se s sed ,h ep r opo sed t h a tt h eg r ea t e rn umb e r o f l i ne s i n f ema le s i n d ica ted p r e f e r en t ia l t r ea tmen t o f m a le s f r om i n f ancy . Women , h e c onc luded , w e r e " t he u nde rnour i shed d r udge s o f t h e i rm enfo lk " ( 1 967 :398 ) . A l though s u b sequen t r e sea r ch ( e .g . G a rn a nd B raun s t e in , 1 986 ; M en s fo r th , 1 9 81 ; Mu r ch i son e ta l , 1 9 84 ) h a s c a s t d oub t u pon t h e v a l id i ty o f H a r r i s l i ne s a s m ea su r e so fh ea l th , We l l s 'r e cogn i t ion o ft h en e ed t o m ove a way f r om t r ad i t iona l a pp roa che s w a s p i one e r ing . C once rn t o r e vea l t h e a n th ropo log i ca l r e le v ance o f o s seou s r ema in s h a s r e su l t ed i n t h e u t i l i sa t ion o f s k e le ta l a nd d en ta l f e a tu r e s w ho se s i gn if i cance i n t e rms o f h ea l th s t a tu s a r e n o t c l ea r ly u nd e r s tood . H a r r i s l i ne s a r e a c l ea r e xamp le ; d en ta l e name l h ypop la s ia ( a d e f ec t p r oduced b y as l ow ing o r c e s sa t ion o f e name l p r oduc t ion d u r ing t h e i n i t ia l s t age o f e name l f o rma t ion ) i sa no the r . E thnog raph ic s t ud ie s a r e e s sen t ia l t o o u r g r ea t e r u nde r s tand ing o f s u ch

w ho le . I tm ay a l so b e ac on sequence o f t h e p ub l i ca t ion o fn ew m ode s o fs k e l e ta l a na ly se s i n s p e c ia l i s t j o u rna l s w i th w h ich a r chaeo log i s t s a r e u na cqua in t ed . Y e t o ne m o re r e a son m ay b e t h e g ene ra l t e nden cy t od e s c r ibe a l s k e le ta l a na ly se s a s " p a la eopa tho logy " . T h i s i s t o o n a r row a d e s c r ip t ion ; i tg i ve s af a l se i mp re s s ion o fw ha ti s , i n f a c t , ab r oad f i e ld o f s t udy . I n deed , a lmo s t t wen ty y ea r s a go , B ro thwe l l ( 1 972 ) e mpha s i sed t h em ino rr o le w h ich p a laeopa tho logy p l ay si nt h e r e con s t ruc t ion o ft h eh ea l th o fp a s tp opu la t ion s . Mo r er e cen t ly , c on s ide rab le e f fo r t s h ave b een m ad e t o p r omo te t h e m u l t i -f a ce ted n a tu r e o f b u r ia l a r chaeo logy i n g ene ra l ( Rob e r t s e t a l , 1 9 89 ) a nd s k e le ta l a na ly s i s i np a r t i cu la r . I n 1 9 87 , f o rt h e f i r s t t i me , as e s s ion d ed i ca t ed t ot h e s t udy o fh uman s k e le ta l r ema in s w a s o r gan i sed a sp a r t o f t h e T heo re t i ca l A r cha eo logy G roup c on f e r en ce , a nd w hen i n t h e f o l low ing y ea r T .A .G . c ame t o S hef f ie ld , w e w e lcomed t h e o ppo r tun i ty t o b u i ld u pon t h i s p r e ceden t . T he t h eme o fo u rs e s s ion w a s i nd e l ibe r a t ec on t r a s tt o t h a to ft h e p r ev iou s y ea r ; a l though w e a pp laud ed t h e i n c lu s ion o f s k e le ta l s t ud ie s i n at h eo r e t ica l c on f e r en ce , w e f e l t t h a t t h e t o ne o f t h e p ape r s , w i th t h e e xcep t ion o f t h a t b y H odge s , w a s s t i l l t h a t o f t r ad i t iona l p a laeopa tho logy , a nd w e b e l ieved t h a t t h e i n te r e s t o fa r chaeo log i s t s m igh t b e s t imu la ted b y a s e r ie s o f p ape r s w h ich d emon s t ra ted t h e " b io cu l tu ra l " a pp roa ch t o s k e le ta l a na ly se s . I n de ed , w e b e l ieve t h i s t ob e t r ue n o tj u s t f o r o u r c on f e r ence s e s s ion , b u t a l so f o rE u rop ean s k e le ta ls t ud ie si ng ene ra l . T he " b iocu l t tn-a l " a pp roa ch r e jec t s t h e t r ad i t i ona l c l in ica l m ode o f s k e le ta l s t udy w h i ch f o cu se s o n t h e i n d iv idua l , a im ing i n s tead a t c omp rehen s ion o f t h e p opu la t ion t o w h i ch t h a t i n d iv idua l b e longed . T he t wo a pp roa che sa r ed i s t i ngu i shed f u r the r i nt h a t r a the r t h an e l u c ida t ing t h e n a tu ra l h i s to ry o fs p e c i f ic d i sea se , t h eb i o cu l t u ra l a pp ro ach i sd i re c t ed t oa n swe r ing q ue s t ion s a bou t t h e g ene ra l h ea l th o f a p opu la t ion , f r equen t ly t h rough t h e a na ly s i s o f ar a nge o f s k e le ta l a nd d en ta l f e a tu r e s , w h ich a r e o f t en r e f e r red t oa s" i nd ica to r so fs t r e s s " .T he se i n c lude t h e a s se s smen t o f a du l t s t a tu re a nd o f s ub -adu l t g r ow th r a te s ; d en ta l c r owd ing ; d e f e c t s o f t h e d en ta l e name l ; o s teopo ro s i s a nd s k e le ta l l e s ion s i n d i ca t ive o f a naem ia . T he i r i n te rp r e ta t ion w a s i n i t ia ted i n r e spon se t o c r i t ic i sm s b y " n ew " a n th ropo log is t s o f t r ad i t iona l s k e le ta l s t ud ie s ( Bu ik s t ra a nd C ook , 1 9 80 ) . H ea l th s t a tu s i nf l u e n ce s l i f e e xpe c tancy , r e p roduc t ive p o t en t ia l a nd t h e c apac i ty f o r l a bou r ( s ee G oodman , t h i s v o lume ) , s o a s B ro thwe l l ( 1 972 :359 ) h a s r i gh t ly s t a ted , " ac on s ide ra t ion o fc ommun i ty h ea l th h a s av e ry r e a l r e levance t ot h e p r ope rs t udy o fe a r ly s o c ie t ie s " . We d o n o t s ugge s t t h a t t h e i d en t i f i ca t i on o f s p e c i f ic d i sea se s h a s n o p l a ce i n t h e r e con s t ruc t ion o f as o c ie ty ' s h ea l th ; t h e p ape r s i n t h i sv o lume b yE i senbe rg ,a nd b yM e i lke john a nd Z ve leb i la r et e s t imony t oi t sv a lue i nt h i sr e spe c t . N o r , i n deed , s hou ld d i sea se a nd h e a l th b e

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a pp roache s i n t h e a r chaeo log ica l c ommun i ty a t l a rge . Wha t , t h en , a r e t h e m a in f e a tu r e s o f t h e b i o cu l tu ra l a pp roa ch ? I n e s sence , t h e b i o cu l tu ra l a pp roach c an b e s e en a s a n e l abo ra t ion o f c oncep t s a nd r e la t ion sh ip s d eve loped i n c u l tu ra l e co logy a nd a pp l ied w i th in t h e f r amewo rk o ft h e e vo lu t iona ry t h eo ry . I n m uch o f t h e e a r l ie r w o rk s t r iv ing t o e l uc ida t e t h e d ynam i c r e la t ion sh ip s b e tween p opu la t ion , c u l tu re a nd t h e e nv i ronmen t , t h e c ond i t ion o f t h e a c to r s -i n d iv idua l s , s o c ia l g r oup s a nd p opu la t ion s -h a s b een n eg le c t ed , e i the rd ue t ot h e l a ck o fp a rad igma t i c a wa r ene s s , o r b ecau se o f p oo r e v idence . I n t r y ing t o u nde r s tand t h e r e la t ion sh ip b e tween p opu la t ion a nd c u l tu r e ,t h ea im o ft h eb i o cu l tu ra la pp roa ch i s t wo fo ld : ( 1 ) t oa s se s s t h e b i o log i ca l c ond i t ion o f h uman p opu la t ion s a nd i t s c on sequen ce s f o r t h e b i o log ica la nd c u l tu ra lr e p roduc t ion o ft h es o c ie ty a nd ( 2 ) t oc on s ide rt h e s e lec t ive e f f ec t so fc u l tu re o n t h e p opu la t ion u nde r s t udy a nd i t s s u rv iva l . U n l ik e s o c iob io logy , w he re g ene t ic i n te r e s t s a r e s e en a s p i vo ta l i ns h ap ing t h e b a s i c s t ruc tu r e o f s o c ie ty , t h e b io cu l tu ra l a pp roa ch p l a ce s e mpha s i s o n t h e i n teg ra t ion a nd i n te r re la t edne s s o f b i o log ica l a nd c u l tu ra l s y s tems , w i thou t a c co rd ing c au sa lp r io r i ty t oe i the r . I np u r su ing t h i s a pp roa ch , w e h ave t oo u t l ine af ew k ey c oncep t s a nd r e la t ion sh ip s , d e sp i te a n i n ev i tab le d i ve rgence o fv i ew sh e ld a bou ts ome o f t h ec omp lex p henomena d i scu s sed b r ief ly b e low : 1 ) T he h uman o r gan i sm , a nd i t s s u rv iv ing r ema in s , i s c omp rehended a s a d ynam ic , h i s to r ica l a nd a dap t ive s y s tem , r ef lec t ing t h e i n te ra c t ion b e tween t h e b i o log ica l o rgan i sm a nd i t s e nv i ronmen t d u r ing t h e i n d iv idua l ' s l i f e t ime . H ea l th i s t h e b i o log ica l a nd p sycho log i ca l c ond i t ion o f t h e i n d iv idua l , as t a te w h i ch c an , a t l e a s t i n t h eo ry , b e a s se s sed b y t h e i n c idence o f i n su l t s a nd o t he r a bno rma l s ymp tom s . T he t wo m a jo r p rob lem s w h i ch a r i se e a r ly i n s u ch a s se s smen t a r e t h e m u l t ip le c au se s w h ich c an r e su l t i n t h e s ame t y pe o f i n su l t , a nd t h e h i e ra r ch ica l o r de r i nw h i ch i n su l t s a r e e xp re s sed , t h e s k e le ton b e ing o ne o f t h e l a s t s y s tems t ob e a f f e c t ed . T he f i r s tp r ob lem m ean s t h a t as ymp tom i sn o tu sua l ly r e la ted t o as i ng le c au se o rs t r e s so r : t h i si sw e l li l u s t ra ted b yD obney a nd G oodman i n t h e i rp ap e ro ne name lh ypop la s ia ,a nd j u s t if ie st h e n eed t ou s e m u l t ip le s t r e s s i n d ica to r s ( i nc lud ing a r cha eo log ica l e v idence ) i n e va lua t ing t h e p o ten t ia l c au se s o f s t r e s s . T he s e cond p rob lem m ean s t h a t s ome i n su l t s , i fa cu te a nd /o r s y s tem s pe c i f ic , m ay n o t l e ave a ny e v idence o f as ke le ta l r e spon se ( s ee G oodman , t h i s v o lume ) . T he i n d iv idua l m ay d i e b e fo r e t h e s k e le ta l r e spon se d eve lop s , o r t h e i n su l t m ay b e r e cove r ed f r om w i thou tl e av ing a ny t r a ce s . H ea l th i s e va lua ted b y c u l tu ra l p e r cep t ion s a nd p a rame te r s : i l -hea l th a nd d i sea se a r e c u l tu ra l ly d ef ined p henomena . I n t h i s n on b i o log ica l s e n se , t h e d ef in i t ion o f h ea l th c an r a nge f r om o ne o f p hy s i ca l a nd m en ta l w e l l -

l e s ion s a nd t h u s t o mo r e a c cu ra t e r e con s t ruc t ion s o ft h eh ea l th o fp a s ts o c ie t ie s . I tw i l lb ea ppa r en tf r om t h i sv o lume t h a tt h e r e i sn o c onsen su s o f o p in ion b e tween r e sea r che r s w i th r e ga rd t ot h e m ea su re s o fh ea l th w h i ch t h ey u t i l i se . S tua r t - Ma cadam , f o r e xamp le , r e i te ra te s h e r p r ev iou s ly ( 1 982 ; 1 985 ) e xp r e s sed v i ew t h a t p o ro t ic h yp e ro s to s is r ef le c t sa naem ia e xpe r ienced i nc h i ldhood ; c onve r se ly ,E i senb e rg r e po r t sa c t ive l e s ion s i na du l tc r an ia . T he d eba te c once rn ing t h e a ge a tw h i ch a naem ic e p i sod e s o ccu r r ed h i nge s o n t h e d i f f e r en t ia t ion o f r emode l led f r om u n re m ode l led l e s ion s ; c u r ren t ly t h e r ea r e n o g ene r a l ly r e cogn i sed s t and a rd s o n w h i ch t o b a se s u ch d i s c r im ina t ion . P a laeodemog raphy a s ab r an ch o f p hy s i ca l a n th ropo logy i sp a r t icu la r ly c on t rov e r s ia l , b e ing v i r tua l ly d i sm is sed b yB o cque t -Appe la nd Ma s se t ( 1 982 ) b u t s t ou t ly d e f ended b y , f o r e xamp le , B u ik s t ra a nd K on ig sbe rg ( 1 985 ) a nd V an G e rven a nd A rme lago s ( 1 983 ) . R e cen t w o rk b y S u chey a nd c o l league s ( Su chey & K a tz , 1 986 ; S uchey e t a l , 1 986 ; S uchey a a l , 1 988 ) o nt h ee s t ima t ion o f a ge -a t -dea th t h rough mo rpho log ica l c hange s o f t h e p ub ic s ymphy sea l f a ce h a s r e v ea led m a jo r d i sc r epanc ie s i n s im i la r , f r equen t ly -u sed t e chn ique s , a nd t h e se d i s c r epanc ie s h ave b een c on f i rmed b y t h e s t udy o f i n d iv idua l s o f k nown a ge -a t -dea th f r om C h r i s t C hu r ch , S p i t a l f ie ld s , ( Cox , p e r s . c omm . ) . T he se r e ve la t ion s h ave s e r iou s i mp l ica t ion s f o r s k e le ta l a na ly se s , s i nce e s t ima t ion s o f a ge -a t -dea th a r e f u ndamen ta l t o a ny a t temp t t o r e con s t r u c t t h e h ea l th o f p a s t s o c ie t ie s . N o r a r e t h e u nce r ta in t ie s o f a ge ing p ecu l ia r t o t h e a du l t s e gmen t o f a s k e le ta l a s semb lage .

The B iocu l tura l Approach i nt h is V o lume I n c ompa r i son t o t r ad i t iona l a pp roa che s t o h uman s k e le ta l r ema in s , t h e b i o cu l tu ra l a pp roa ch i s an ew a r ea , a nd a s w i th a ny n ew d i s c ip l ine , o ve r s imp l i f ica t ion a nd v a r ia t ion s i n i n te rp r e ta t ion a r e t o b e e xpe c t ed , p a r t i cu la r ly i n o ne w h ich i n co rpo r a te s a s m any u nknown v a r iab les a sd oe s a r chaeo logy . T he r e cogn i t ion o f t h e se p rob lems s hou ld b e v i ewed i n ap o s i t ive l i gh t s i nce i ts t imu la te s f u r the r r e sea r ch , w h i le i n t roduc ing r e a l i sm t os k e le ta l s t ud ie s . I nc on t r a s t t oN o r th Ame r ica , t h e b i o cu l tu ra l a pp roa ch i sy e t t o b ecome e s tab l i shed i n E u rope , b u t a l though p opu la t ion a na ly se s a r e l a ck ing , w i th t h e e xcep t ion o f S t . H e len s -on the - Wa l l s ( G r aue r , 1 989 ) , t h eo re t ica l a nd m e thodo log ica l s t ud ie s o f m ea su re s o f h ea l th a r e mo r e a bundan t ( e .g . B u l l ion , 1 987 ; B u sh , 1 990 ; D obney , 1 983 , f o r thcom ing ; S t ua r t - Ma cad am , 1 982 ) . T he p u rpo se o f t h e T .A .G . c on f e r en ce s e s s ion w h ich w e o rgan i sed i n 1 988 , a nd i t s t r an s la t ion , w i th a dd i t iona l p ape r s , i n to t h i s v o lume , w a s t o i l u s t ra te r e sea r ch i n sp i red b yb i o cu l tu r a lc oncep t s o n b o th s i des o f t h e A t l an t ic a nd p romo t e s u ch

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f u ndamen t a l f o r ce s i n t h e c on t inu ing s t o ry o f h uman e x i s t ence . E ven am ode ra te ly c omp rehen s ive s u rvey o f t h e se c oncep t s i sb eyond t h e s cop e o ft h e p r e sen t v o lume ,a nd d e se rve s at ome i ni t so wn r i gh t . B u t s i n ce t h e s e c oncep t s a r e s o g e rmane t o t h e a pp roa ch a dop ted i nt h i s v o lume , a n o pe r a t iona l a nd h eu r i s t i cc ha ra c te r isa t ion i so f f e r ed b e low : a ) T he r e a r e d ynam ic l i nk s b e tween t h e n a tu r a l e nv i ronmen t , t h e s o c ia l e nv i ronmen t a nd h uman b i o logy . C hange o ccu r s t h rough s e lec t ion ( whe the r c u l tu r a l o r n a tu ra l ) a c t ing o n t h e v a r iab i l i ty i n h uma n b ehav iou r a nd h uma n b i o logy , s e lec t ing f o r t h o se t r a i t s w h ich a r e a dap t ive i n t h e r e la t ion sh ip t o t h e s o c ia l a nd n a tu r a l e nv i ronmen t o f t h e t i me ( s ee a l so t h e c on t r ibu t ion s t o t h i s v o lume b y E i senbe rg a nd G oodma n ) . b ) A dap t ivene s s c an b e d e f ined i n t e rms o f i n c lu s ive D a rw in ian f i t ne s s : t h e g r ea te r c hance s o f s ome i n d iv idua l s t o s u rv ive a nd r e p roduce , r e la t ive t o o t he r s , i n a g i ven s o c ia l a nd e nv i ronmen t a l s i tua t ion ( b u t s e e H y land a nd S cu t t ,t h i sv o lume ) . c )H ea l th a nd d i sea se a r e c ond i t ion s a f f e c t ing t h e c hance s o f i n d iv idua l s t o s u rv ive a nd r e p rodu ce , a nd , b y e x ten s ion , t h e r e p roduc t ion o f t h e s o c ia l e nv i ronmen t . H ea l th a nd d i sea s e , t h en , a r em a jo re l emen t so fs e le c t ion . d ) A t t h e s ame t i me , h ea l th a nd d i sea se a r e p r oduc t s o f h uman b i o log ica l v a r iab i l i ty ( i n t h e g eno typ e a nd t h e p heno type ) , a nd t h e s e le c t ive p r e s su r e s i mpo sed o n i tb y t h e s o c ia l a nd n a tu ra l e nv i ronmen t . B ehav iou r w h ich f a i l s t o r e so lv e t h e se p r e s su r e s t o t h e b ene f i t o f t h e i n d iv idua l " c an r e su l t i n i mpa i r ed p hy s ica l a nd p sy cho log ica l w e l l -b e ing , i n d i sea se a nd u l t ima te ly i n d ea th " ( Cox , 1 983 :255 ) . T he c ond i t ion o f h e a l th , t h en , c an b e s e en a l so a s a r ef le c t ion o f t h e h uman b i o log ica l a dap ta t ion t o i t se nv i ronmen t( s e eG oodman ,t h i s v o lume ) . An umbe ro fp ape r sa dd r e s st h e se i s sue si nt h e p r e sen t v o lume . E i senbe rg , i n h e r s t udy o f t h e M idd le C umb e r land g roup s i nT enne see , c on s id e r s t h e i mp l i ca t ion s o f d i e ta ry s t re s s a s w e l l a s s o c ia l s t r e s s ( s ee a l so J ohn son , 1 982 , f o r t h e n o t ion o f s c a la r s t r es s ) , b o th p roduc t s o f t h e s o c ia l e nv ironmen t a s w e l l a s n a tu ra l c ond i t ion s . A r chaeo log ica l ,c u l tu ra ld a ta p l ay ac r uc ia lr o le i n i d en t i fy ing t h e c on s t ra in t s i mpo sed b y t h e s o c ia l e nv i ronmen t , a nd E i senb e rg ' s i n t e rp r e ta t ion c l ea r ly w ou ld n o t b e a s c omp le te h ad e i the r t h e b i o log ica lo rc u l tu r a ld a ta b een o m i t ted . G oodman p r e sen t s am ode l i nw h ich h ea l th i s ac en t ra l f e a tu r e o f t h e a dap t ive p ro ce s s , a nd c on s ide r s t h e p o s i t ion o f h ea l th i np romo t ing o r h i nde r ing c u l tu ra l a nd s o c ia l r e p roduc t ion . T he c on t r ibu t ion so fC a t taneo a nd o fH y land a nd S cu t t a dd r e s s t h e q ue s t ion o f g ene t i c a s o ppo s ed t o e nv i ronmen ta l f a c to r s i n t h e h ea l th s t a tu s o f i n d iv idua l sa nd p opu la t ion s . H y land a nd S cu t t g o b eyond t h i s c on t rove r sy a nd p o s i t t h e c on t r a d i c t ion b e twe en i n d iv idua l s u rv iva l a nd g ene t ic

b e ing , w i th a ny th ing e l se b e ing i l -hea l th , t ot h e c l in ica l p r e sence o f d i sea se i n d ica to r s o r o f t h e b i o log ica l r e spon se t oi t( s ee H y land a nd S cu t t ' s d i s cu s s ion o ft h e " med ica l mode l " ) w i th a ny th ing o t he r c on s ide r ed h ea l thy . T h i s s e rve s t oi l u s t ra te t h a t t h e c oncep t o f h ea l th i t se l f i s ap rob lema t ic a nd c u l tu r e spe c i f ic n o t ion , a nd p o in t s t o t h e d i f f e rence s b e tween c u l tu ra l a nd m ed i ca l / c l in ica l p e r cep t ion s o fh ea l th ( s ee B ush , t h i s v o lume ) . A s s t uden t s o f p a s t s o c ie t ie s , w e a r e i n te r e s ted i n b o th p e r cep t ion s o fh ea l th , b u to n ly r a r e ly a nd b y r e f e r ence t ot h e n o t too -d i s tan tp a s ta r e w ea b le t o u nde r s tand t h e c u l tu ra l d e f in i t ion . T h i s l e ave s u s w i th t h e c l in ica l a s se s smen t o fh ea l th a st h e o n ly c r o s s -cu l tu ra l w ay o fe va lua t ing t h e h ea l th s t a tu s o f p a s t p opu la t ion s f r om t h e a r chaeo log ica l r e co rd , y e t a s B u sh d e s c r ibe s i n t h is v o lume , c l in ica l d e f in i t ion s o f h ea l th c anno t p rov ide a c omp rehen s ive f r amewo rk f o r d e s c r ib ing t h e c ond i t ion . 2 ) S i nce h uman s a r e s o c ia l c r ea tu r e s w ho s p end m o s t o f t h e i r l i ve s i n t h e c ompany o f o t he r s , t h e h ea l th o f i n d iv idua l s c anno t b e a s se s sed f u l ly w i thou t e va lua t ing t h e h ea l th o f o t he r i n d iv idua ls w i th in a c ommun i ty . E qua l ly , s i nce h umans d epend f o rs u rv iva lo nc o -ope r a t ion o rc ompe t i t ion w i th o t he r s a nd s i n ce t h e p a s sage o f s o c ia l t r ad i t ion s i s s ub jec t t o c on t ro l a t h ou seho ld , c ommun i ty a nd b r oad e r p opu la t ion l e ve l s , o n ly t h e s t udy o f h ea l th a nd d emog r aph i c p a rame te r s a tt h e se l e ve l s c an a n swe r q ue s t ion s a bou t t h e i nf luence o f h ea l th o n b i o log i ca l a nd s o c ia l r e p roduc t ion . W i th in t h e b i o cu l tu ra l a pp roa ch , t h e re fo re , t h ee mpha s i s i sp l aced o n t h e h ea l th c ond i t ion o fc ommun i t ie s a nd p opu la t ion s , a sw e l l a si n d iv idua l s . T he c oncep t o fc ommun i ty h ea l th a nd i t s a pp l i ca t ion s a r e d eve loped f u r the r b yE i senbe rg i nt h is v o lume . 3 ) T he k ey o b jec t ive o f t h e b i o cu l tu ra l a pp ro ach i st ou nde r s tand t h er o le o fh ea l th i nt h e d ynam ic i n te r ac t ion b e tween t h e e nv i ronmen t , h uman p opu la t ions a nd c u l tu r e . S uch a n o b jec t ive a dd r e s se s k ey c oncep t s i n t h e s t udy o f h uman c u l tu ra l a nd b i o log i ca l e vo lu t ion , s u ch a st h o se o f a dap ta t ion a nd f i t ne s s . A sG oodman e ta ln o ted i n a ne a r l ie rp ub l ica t ion : " . . s t re s s i s ap o ten t ia l ly s i gn if i can t f o r ce f o r c hange i nf i t ne s s o f c u l tu ra l a nd b i o log ica l s y s tem s , at o p i c o fc en t ra l c on c e rn a mong a n th ropo log i s t s " ( Goodman e t a l , 1 980 :516 ) . T he se c oncep t s a nd r e la t ion sh ip s a r e n o t e a sy t od ef ine a nd d i f f i cu l t t oi l u s t ra te w i thou t g ro s s g ene ra l i sa t ion s . I n a r chaeo logy , w e h ave o n ly v ague a nd d i sco rdan t n o t ions o ft h e m ean ing a nd r o le o f s u ch p ro ce s se s a s n a tu ra l s e lec t ion a s o ppo sed t oc u l tu ra l s e lec t ion a nd a dap ta t ion , a nd o f s u ch s t a te s a s f i t ne s s ( e .g . D unne 1 1 , 1 980 ; I n go Id , 1 986 ; M i then , 1 989 ; R indo s , 1 984 , 1 985 ; R edd ing , 1 988 , e t c . ) . A nd y e t t h e b i o cu l tu r a l a pp roa ch a im s a t e l uc ida t ing t h e r o le o f t h e se

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i se ncoun te r ed . S ome i n su l t s w i l l n o t a f f ec t t h e s k e le ta l a nd d en ta l s y s tems , w h i le o t he r s m ay b e s o a cu te t h a t t h e i n d iv idua l d i e s b efo re t h e se s y s t em s a r e i n vo lved . I nc onne c t ion w i th t h i s , t h e p o in tm ade f o r ce fu l ly b yG oodman i nt h i sv o lume i sw o r th r ememb e r ing : m a rke r s o fi n su l t a r es i gn s o f as t rugg le t oa dap t . I th a s b een a s sumed i nt h e p a s t t h a t ag r ea te r n umbe r o f m a rke r s o f i n su l t m ean s ag r ea te rd eg r ee o fi l -hea l th ,b u t i tm ay b e t h a t t h e o ppo s i t e i st r i te , f o r n o t o n ly a r e s u ch m a rke r s t e s t imony o f a n e ncoun te r w i th s t r e s s , b u t a l so o f r a l ly ing f r om i t . T h i s i s w e l l d o cumen ted b y t h e e xamp le o f s e ve r e i n f ec t ion : s u ch a c ond i t ion w ou ld m ake h eavy d emand s u pon t h e i mmune s y s tem , w i th a c on sequen t l owe r ing o f c hance s o f s u rv iva l ( La l lo e t a l , 1 9 78 ) , y e t p a radox i ca l ly , s i nce b one i sg ene ra l ly t h e l a s t s y s tem o f t h e b ody t o b e a f f e c ted b y i n fe c t ion , t h e i mmune s y s tem h a s t o b e a b le t o r e spond s u f f i c ien t ly f o rt h es k e le ton t ob ea f f ec ted ( O r tne r , 1 988 ) . Af u r the r m a jo r p o in t t oc on s ide r c once rn ing t h e f r equency o f m a rke r s o f i n su l t i s t h a t t h e r e spon se t ot h e " s t re s s " w h ich p roduced t h em i s n o t u n i fo rm e i the r w i th in o r b e tween i n d iv idua l s ( s ee B u sh , t h i s v o lume ) . T hu s t h e r e m ay b e t empo ra l v a r ia t ion i n r e s i s tance , s t r e s s b e ing e xpe r ien ced o n ly w hen r e s i s tance i sl owe r ed . T h i s l e ad s t o ap r ob lem o fi n te rp r e ta t ion : a b sence o fa m a rke r o fi n su l t m ay b e i n d ica t ive o f h e igh tened r e s i s tance , b u t c onve r se ly i tm ay m ean f a i lu re t o e n coun t e r as t re s s invok ing s i tua t ion . Mo r eove r , v a r ia t ion o fr e spon se b e tween i n d iv idua ls m ay b e ac i r cums tance o fg ene t ic f a c to r s , a ge a nd g ende r ( Goodman e ta l , 1 984 ) . T he r e la t ive f r equency o f s k e le ta l m a rke r s o fi n su l t i nm a le s a nd f ema le s i s r o u t ine ly a s se s sed i na r cha eo log ica l a s semb lage s a nd f r equen t ly i n te rp r e ted i n c u l tu r a l t e rm s ; We l l s ' u s e o f H a r r i s l i ne s , t o t a ke j u s t o ne e xamp le , w a s r e f e r r ed t oe a r l ie r . I n t h e f a ce o f s u ch c u l tu ra l i n te rp re ta t ion s o f m a rke r s o f i n su l t , i ti sn o t ewo r thy t h a t w omen m ay b e h ypoac t ive w i th r e ga rd t ot h e i r s ympa the t ic -ad r enomedu l la ry r e spon se ( Cox , 1 978 ) , f o rt h i s m ay w e l l i nf luence t h e f o rma t ion o f m a rke r s i nw omen . S o t o o m ay t h e g r ea t e r r e s i s tance o f g i r l s t h an b oys t o t h e c on sequen ce s o fi l ne s sa nd m a lnu t r i t ion ( Tanne r , 1 9 77 ) . V a r ia t ion i nt h e f r equenc ie s o f m a rk e r s o f i n su l t i n w omen a s c ompa r ed t om en m ay b e a r ef l ec t ion , n o t o f c u l tu ra l p r ac t ice s , b u t o f d i f f e r ing p hy s io log ica la c t iv i ty . A s t h i s s ho r t i n t roduc t ion s how s , t h e s t udy o f h ea l th i np a s ts o c ie t ie s , f r om t h er e con s t ruc t ion o f b a s ic p a rame t e r s t o t h e a s se s smen t o f w ide r i mp l ica t ion s , i s av e ry c omp lex f i e ld i nt h e e a r ly s t age so fp r ac t i ce . I no u rv i ew , h oweve r , h ea l th a s a m ea su r e o f f i t ne s s , a nd a s a c ond i t ion i nf luenc ing s o c ia l a nd b i o log ica l r e p roduc t ion , i s c en t ra l t oo u r u nde r s tand ing o f h uman b i o log ica l a nd c u l tu ra l d eve lopmen t . I t i s o n ly a t t h e i n te r f a ce o fa r cha eo logy a nd c u l tu ra la nd p hy s ica l a n th ropo logy , w i th in t h e f r amewo rk o f t h e

e vo lu t ion . A dop t ing t r ou t a s a n a na logue , t h ey d e f ine b i o log ica la dap ta t ion a st h ec on t inua t ion o f t h e g ene p oo l , a nd e mpha s i se r e p roduc t ion a st h e k ey e l emen t i nt h e d e f in i t ion o ff i t ne s s . T h i s m ay b e a t t h e e xp en se o f s u rv iva l : s o l o ng a s t h e i n d iv idua l s s u rv ive l o ng e nough t op a s s o n t h e i r g ene t ic i n he r i tance , t h ey m ay b e d eemed f i t . T h i s i st h ough t -p rovok ing , b u t a s H y land a nd S cu t t p o in t o u t , t h e i n c rea sed l o ngev i ty o f modem h uman s , w e l l b eyond t h e i r p r ime r e p rodu c t ive y ea r s , m ay b e s e en a s ac u l tu ra l a dap ta t ion t o e duca te a nd p a s s o n s o c i a l t r ad i t ion s f r om o ne g ene ra t ion t o a no the r , t h e r eby i n c rea s ing t h e s u rv iva l a nd r e p roduc t ive c hance s o f t h e o f f sp r ing . N a tu r a l s e le c t ion , t h e r ef o r e , w ou ld s e lec t f o r c u l tu re a nd t h e g r ow th o f s o c ia l t r ad i t ions , s o l o ng a s i t c onf e r s s e l ec t ive a dvan tage o n c u l tu re -cons c iou s g roup s i n r e la t ion t o o t her s ( o u r e mpha s i s ) . T he i mp l ica t ion s o ft h e r e la t ive n a tu r e o ft h e s u r v iva l v a lue o fc u l tu re a r ey e tt ob ee xp lo r ed . H y land a nd S cu t t ' sp ap e rc a s t s ar e la t ive v a lue o n p a laeodemog raph ic m ea su r e s s u ch a st h e l i f e e xpec tancy , s u rv ivo r sh ip , m o rb id i ty a nd m o r ta l i ty o f ap opu la t ion a s t h e u l t ima te a nd c r o s s -cu l tu ra l m ea su r e so fa dap t ive f i t ne s s . 4 ) A t am e thodo log i ca l l e ve l , t wo m a jo r p r ob lem a r ea s e x i s t . F i r s t h ow f a r i s t h e b u r ia l /mo r tua ry /mo r ta l i ty s amp le r e p re sen ta t ive o f t h e c ond i t ion s i n t h e l i v ing p opu la t ion f r om w h ich i t w a s t a ken , a nd s e cond , h ow r e p r e sen ta t ive o f i n d iv idua l h ea l th s t a tu s a r e s k e le ta l a nd d en ta l m a rke r s o f i n su l t ? T he v e ry f a c t t h a t a r chaeo log ica l s k e le ta l a s semb lage s a r e m o r ta l i ty s amp le s m ea n s t h a t t h ey a r e u n l ik e ly t o b e ad i re c tr e f lec t ion o f al i v ing p opu la t ion ( Cook , 1 981 ) . We h ave t ob e a wa r eo f ar a nge o fb i a s ing f a c to r s : a ) s o c ia l t r ad i t ion s a nd i d eo logy m ed ia te b e tween t h e l i v ing a nd t h e d ead a nd a c t a s a s e lec t ive c ode a s t o w he r e a nd h ow p eop le a r e b u r ied ,a nd w ho i sa nd w ho i sn o tt r ea ted t h u s . b ) p ro ce s se s o f d i f f e ren t ia l p r e se rva t ion a nd a l te r a t ion a c tu pon t h eb u r ied h uman r ema in s . c ) a n umbe r o f f a c to r s e n su r e t h a t t h e f r equency o f m a rke r s o f i n su l t i n as k e le ta l p opu la t ion w i l l n o t r ef le c t t h a t o f t h e l i v ing p opu la t ion f r om w h i ch i tw a s d e r ived . T he i r f r equency w i l l b e a f f e c t ed b y t h e i n te r ac t ion o f ( 1 ) t h e a ge s t ruc tu re o ft h e i n d i ca to r i nt h e l i v ing p opu la t ion a nd a ge spec i f i c d ea th r a te s a nd ( 2 ) c au se s o fd ea th a nd t h e f r equency o ft h ei n d ica to r ( Cook , 1 981 ) . I ti sc ommon p r a c t ice t oc ompa r e s k e le ta l a s semb lage s w h i ch d i f f e rc h rono log ica l ly o rg eog raph ica l ly ,y e tC ook ( i b id . ) h a ss hown t h a t t h e d eg ree o f g r ow th d i s tu rbance , a nd b y e x ten s ion , t h e f r equency o f m a rk e r s o f i n su l t , i s u n l ike ly t o b e c ompa rab le b e tween t wo d i s c r e te p opu la t ion s i n w h ich t h e c au se s o f d ea th a r e d i f f e ren t . T he d eg r ee t o w h ich t h e h ea l th s t a tu s o f a n i n d iv idua l i s r ef le c ted s k e l e ta l ly d epend s t o a c e r ta in e x ten t u pon t h e n a tu r e o ft h e i n su l t w h ich

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e d s . C . R enf r ew , M . R ow land s & B . S eag rave s . p p . 3 89 -4 1 .O r lando :A cadem ic P r e s s . Men sfo r th , R .P . ( 1 981) . G row th v e loc i t y a nd c hond rob las t i c s t ab i l i ty a s m a jo rf ac to rs i nf l uenc ing t h ep a thogene s i s a nd e p idem io log ica l d i s t r ibu t ion o f g row th a r res t l i ne s . P ap e r p r e sen ted a t F i f t ie th A nnua l Mee t ing o f t h e Ame r ican A s soc ia t ion o f P hy s ica lA n th ropo log i s t s ,D e t ro i t . M i then , S . ( 1989 ) . E vo lu t iona ry t h eo ry a nd p os t p roce s sua la r chaeo logy .A n t iqu i ty 6 3 :483 -94 . Mood ie , R .L . ( 1 923 ) . P a laeopa tho logy : a n i n t ro d uc t ion t ot h e s t ud y o fa nc ien t e v idences o fd i sea se . U rbana ,I l ino i s : U n ive r s i ty o fI l ino i sP re s s . Mu rch i son , M .A . , Ow s ley , D .W . & R iope l le , A . J . ( 1 984 ) . T r an sve r se l i ne f o rma t ion i n p ro te ind ep r ived r h e su sm onkey s . H um .B io l . 5 6 :173 -182 . O r tne r , D . J .( 1 988 ) . T heo re t ica l a nd Me thodo log ica l I s sue s i nP a l eopa tho logy . P ape r p repa red i na dvance f o r p a r t ic ipan t s i n t h e I n te rna t iona l C ong r e s s o f A n th ropo log ica l a nd E thno log ica l S c ience s S ympo s ium o n H uman P a leopa tho logy : C u r ren t S yn the se sa nd F u tu re O p t ion s . R edd ing , R .W . ( 1 988 ) . A g ene ra l e xp lana t ion o f s ub s i s tence c hange : f r om h un t ing a nd g a th e r ing t o f o od p r oduc t ion .J .A n th ropo l .A rchaeo l . : 5 6 -97 . R indo s ,D . ( 1984 ) . T he O r ig in o fA gr icu l tura lS y s tem s : A n E vo lu t iona ry P e r spec t ive . O r lando : A cadem ic P re s s . R indo s , D . ( 1 985 ) . D a rw in ian s e lec t ion , s ymbo l ic v a r ia t ion a nd t h e e vo lu t ion o f c u l tu re . C ur l . A n thropo l .2 6 :65 -88 . R obe r t s , C .A . , L ee ,F .&B in t l i f f , J .( 1989 ) . e d s .B ur ia l A rchaeo logy : C u rren t R e sea rch , Me thod s a nd D eve lopmen t s . B r i t i sh A rchaeo log ica l R epo r t s , B r i t i sh S e r ie s2 11 . R oge r s , A .R . ( 1 988 ) . D oe s b i o logy c ons t ra in c u l tu re? Am .A n th ropo l .9 0 :819 -31 . R oge r s , J . a nd D iepp e , P . ( 1 983 ) . L e s son s f r om P a la eopa tho logy . T he P rac t i t ione r2 27 :1191 -99 . S t ua r t - Macadam , P . ( 1 982 ) . AC o r re la t ive S t udy o fa P a laeopa tho logy o f t h e S ku l l . U npub l i shed P h .D . t h e s i s ,U n ive r s i ty o fC amb r idge . S t ua r t - Macadam , P . ( 1985 ) . P o ro t ic h yp e ro s to s i s : r e p r e sen ta t iv e o f ac h i ldhood c ond i t ion . Am .J .P hys . A n thropo l . 6 :39 1-8 . S uchey , J .M . a nd K a tz , D . ( 1986 ) . S k e l e ta l A ge S t anda rd s D e r ived f rom a n E x tens ive Mu l t i rac ia l S amp le o f Mode rn Ame r ican s . P ap e r p r e sen t ed a tt h e F i f ty F i f th A nnua l Mee t ing o f t h e Ame r ican A s soc ia t ion o f P hy s ica l A n th ropo log i s t s , A lbuque rque ,N ew Mex ico , A p r i l 1 986 . S uchey , J .M . , B rooke s , S .T . & K a tz , D . ( 1 988 ) . I n s t ruc t iona l m a te r ia l s a ccompany ing f ema l e p ub ic s ymphy sea l m od e l s o f t h e S uchey -Brooke s s y s tem . D i s t r ibu ted b yF rance C a s t ing . S uchey , J .M . , W i se ly , D .V . & K a tz , D . ( 1 986 ) . E va lua t ion o ft h eT odd a nd McKe rn -S tewa r tm e thod s f o r a ge ing t h e m a le o sp ub i s . I nF o ren s ic O s teo logy : A dvance si nt h eI d en t if ica t ion o f Human R ema in s . e d . K . R e ich s . p p . 3-67 . S p r ingf ie ld , I l ino i s : C ha r le s C . T homa s . T anne r , J .M . ( 1 977 ) . P hy s ica l a nd p sycho log ica l d eve lopmen t . I nH uman B io logy . 2 nd e d . e d s . G .A . H a r r i son , J . S . We ine r , J .M . T anne r &N .A . B a rn ico t . p p .3 52 -360 .O xf o rd : O xf o rd U n ive r s i ty P re s s . V an G e rven , D .P . a nd A rme lago s , G . J . ( 1 983 ) . " Fa r ewe l l t o P a leodemog raphy? " R umou r s o f i t s d ea th h ave b e en g r ea t ly e xagge ra ted . J .H um . E vo l . 1 2 :353 -60 .

b i o cu l tu ra l a pp roa ch , t h a t t h e s t udy o f h ea l th i n p a s ts o c ie t ie sc an b ef u l ly d ev e loped .

R e ferences A cke rknech t , E .H . ( 1953 ) . P a leopa tho logy . I n A n th ropo logy T oday , e d . A .L . K roebe r . p p . 1 20 -26 . C h icago :U n ive r s i ty o fC h icago P re s s . B o cque t -Appe l ,J P . a nd Ma s se t ,C . ( 1 982 ) .F a r ewe l l t o p a leodemog raphy .J .H um .E vo l . 1 :321-33 . B ro thwel , D .R . ( 1 972 ) . C ommun i ty h ea l th a s af a c to r i nu rban c u l tu ra l e vo lu t ion . I n Man , S e t t lemen t a nd U rban i sm , e d s . P . J . U cko , R . T r ingham a nd G .W . D imb leby .p p . 3 53 -61 .L ondon : D uckwo r th . B u ik s t ra , J . E . a nd C ook , D .C . ( 1 980 ) . P a la eopa tho logy : a n Ame r ican a ccoun t . A nn . R ev . A n thropo l . 9 : 433 4 70 . B u ik s t ra , J . E . a nd K on i sbe rg , L .W . ( 1 985 ) . P a le o de rnog raphy : c r i t ique s a nd c on t rove r s ie s . Am . A n th ropo l .8 7 :316 -33 . B u l l ion , S . ( 1987 ) . I ncremen ta l S t ruc ture s o fE name l a nd t h e ir A pp l ica t ions t oA rcha eo logy . U npub l i shed P h .D .t h e s i s ,U n ive r s i ty o fL anca s te r . B u sh , H .M . ( 1990 ) . T he R ecogn i t ion o fP hys io log ica l S t res s i n H uman S ke l e ta l R ema in s : aC r i t ique o f Me thod a nd T heo ry w i th S pec if ic R ef e rence t o t h e V e r teb ra l C o lumn . U npub l i shed P h .D . t h e s i s , U n ive r s i ty o fS he f f ie ld . C ook , D .C . ( 1981) . Mo r ta l i ty , a ge st ruc tu re a nd s t a tu s i nt h e i n te rp re ta t ion o fs t r e s s i n d ica to r s i np r eh i s to r ic s k e le ton s : ad en ta l e xamp le f r om t h e L owe r I l ino i s V a l ley . I n T he A rchaeo logy o f D ea th . e d s . R . C hapman , I .K inne s & K . R and sbo rg . p p . 1 33 -44 . C amb r idge : C amb r idge U n ive r s i ty P r e s s . C ox , T .( 1978 ) .S t re s s .L ondon : MacM i l lan P r e s sL td . D a s tugue , J . ( 1 980 ) . P o s s ib i l i t ie s , l i m i ta t ion s a nd p ro spec t so fp a leopa tho logy o ft h eh uman s k e le ton .J . H um .E vo l . 9 : 3 -8 . D enne l l , R . W . ( 1979 ) . P reh i s to r ic d i e t a nd n u t r i t ion : s ome f ood f o rt h ough t . Wo r ld A rchaeo l . 1 :121 -35 . D obney , K . ( 1983 ) . E name l H ypop la s ia o fP ig T ee th : i t s R e levance a nd A pp l ica t ion t oZ ooa rchaeo log ica l S t ud ie s . M .Sc .T he s i s ,U n ive r s i ty o fL ondon . D obney , K . ( f o r thcom ing )A nE p id em io log ica lS t udy o f D en ta l E name l H ypop las ia i n B radf o rd S choo l C h i ld ren : i t s R e levance t o A rchaeo log ica l S t ud ie s . P h .D .t h e s i s ,U n ive r s i ty o fB rad f o rd . D unne 1 1 , R .C . ( 1980 ) . E vo lu t iona ry T heo ry a nd A rchaeo logy . I nA dvances i nA rchaeo log ica l Me thod a nd T heory V o l . 3 . e d . M . S ch i f f e r . p p . 3 8 -99 . O r lando :A cad em ic P re s s . G a rn , S .M . a nd B raun s te in , E .M . ( 1 986 ) . C on tr ibu t ion s o fm u l t i imag ing t e chn ique s t oa n u nde r s tand ing o f s t ruc ture sr ep resen ted b yr ad iopaque l i ne sa nd b and s i n t u bu la r b ones . P ape r p r e sen t ed a t5 5 th A nnua l Mee t ing o f Ame r ican A s soc ia t ion o f P hy s ica l A n th ropo log i s t s , A lbuque rque , N ew Mex ico , A p r i l 9 1 2 . G oodman , A .H . , A rme lagos , G .J . &R o se , J . C . ( 1 984 ) . T he c h rono log ica l d i s t r ibu t ion o fe name l h ypop la s ia s f r om p r eh i s to r ic D ick son Mound sp opu la t ion s . Am .J . P hys .A n th ropo l . 6 5 :259 -66 . G raue r , A .G . ( 1989 ) . H ea l th , D i sea se a nd S t a tu s i n Med ieva l Y o rk . U npub l i shed P h .D . t h e s i s , U n ive r s i ty o f Ma s sachu s se t t s , Amhe r s t . I n go ld , T . ( 1 986 ) . E vo lu t ion a nd S oc ia l L i f e C amb r idge : C amb r idge U n ive r s i ty P r e s s . J ohn son , G . ( 1982 ) . O rgan iza t iona l s t ruc tu re a nd s c a la r s t re s s . I n T heo ry a nd E xp lana t ion i nA rchaeo logy .

8

B ro thwe l l a nd A .T . S p r ingf i e ld : T homa s .

Wa shbu rn , S .L . ( 1953 ) . T he s t ra tegy o f p hy s ica l a n th ropo logy . I n A n thropo logy T oday , e d . A .L . K roebe r , p p . 7 14 -27 . C h icago : U n ive r s i ty o fC h icago P re ss . We l l s , C . ( 1961) . An ew a pp roach t o a nc ien t d i sea se . D iscove ry 2 2 :526 -31 . We l l s , C . ( 1964 ) . B one s , B od ie s a nd D i sease . L ondon : T hame sa nd H ud son . We l l s , C . ( 1967 ) . An ew a pp roa ch t op a laeopa tho logy : H a r r i s ' l i ne s . I n D i sea se s i nA n t iqu i ty e d s . D .R .

S and i son .

p p .

3 90 -404 .

A cknow ledgemen t We t h ank D r . Ma rga r e t C ox f o r u npub l i shed d a ta r e la t ing t ot h e s k e le ta l r ema in s f r om C h r i s t C hu r ch ,S p i ta l f ie ld s .

9

2 .

CONCEPTS OF HEALTH AND S TRESS

H e len B ush b een ar e v iva l o ft h e e co log i ca l c oncep to fh ea l th , w i th ar e su rgence o f i n te re s t i nh omeopa thy a nd o t he rh o l i s t ic , " a l t e rna t ive " f o rm so fm ed i c ine . T he e co log ica ld e f in i t ion o fh ea l th ,w h ich s e e s h ea l th a nd d i sea se a ss u cce s s fu la nd u n succe s s fu l a t t emp t s t o a dap t t o s t r e s s f r om i n te rna l a nd e x t e rna l s ou r ce s , u nde r l ie s t h e b i o cu l tu ra l a na ly s i s o f s k e le ta l r ema in s . I te mpha s i se s t h e d ynam ic n a tu r e o f t h e c ond i t ion , s p ec if ica l ly t h e n o t ion o f a d ju s tmen t t o , o r r a l ly ing f r om , u nf avou rab le i nf luence s . F o re xamp le :

I n troduc t ion E co log ica l d e f in i t ion s o fh ea l th e mpha s i se t h e d ynam ic n a tu re o ft h ec ond i t ion , a nd i np a r t icu la r t h ea b i l i ty o ft h ei n d iv idua lt or e cove rf r om i n su l t . I n c r ea s ing ly , a n th ropo log i s t s e va lua t e t h e h ea l th o f p a s t p opu la t ion s , r e p r e sen ted b y s k e le ta l a s semb lage s , t h rough t h e i n te rp r e ta t ion o fd en ta l a nd s ke le ta l f e a tu r e s c au sed b y t h e h uman o rgan ism ' s a dap t ive r e spon se . T he s e m a rke r s a r e c ommon ly k nown a s" i nd i ca to r so fs t r e s s " . T h i s c hap t e r e xp lo r e s m ed i ca l a nd a n th ropo log i ca l c oncep t s o f s t r e s s , a nd p ropo se s t h a t t h e a dop t ion o f S e ly e ' s p hy s io log i ca l s t re s s mod e l b y a n th ropo log i s t s h a s m ean t t h a t p sycho log i ca l i nf luence s o n t h e f o rma t ion o f s t re s s m a rke r s h ave b een i g no r ed , b y a nd l a rge . T he c hap te r p r e sen t s e xamp le s f r om t h e f i e ld s o f m ed ic ine a nd p y scho logy t o d emon s t r a te t h e i mpac t w h ich a dve r se m en ta l s t a te s m ay h ave u pon t h e g r ow th p ro ce s s , a nd i ti sc on c luded t h a t i ti s e s sen t ia l t o k eep i n m ind p sycho log ica l f a c to r sw hen i n te rp r e t ing i n d ica to r so fs t r e s s .

" Hea l th i s a c on t inu ing p rope r ty , p o ten t ia l ly m ea su rab le b y t h e i n d iv idua l ' s a b i l i ty t o r a l ly f r om i n su l t s , w he the r c hem i ca l , p hy s i ca l , i n f ec t iou s , p sy cho log i ca l o r s o c ia l " ( Audy , 1 9 71 :142 ) . " Hea l th i s af u nc t ion o ft h e o r gan i sm ' s a b i l i ty t o a d ju s t t o e nv i ronmen ta l c on s t ra in t s a nd s t re s so r s " ( Goodman , t h i s v o lume ) . " S ta t e s o f h ea l th o r d i sea se a r e t h e e xp r e s s ion o f t h e s u cce s s o r f a i lu r e e xp e r ienced b y t h e o r gan i sm i ni t s e f fo r t s t o r e spond a dap t ive ly t o e nv i ronmen ta l c ha l lenge s " ( Dubo s , 1 965 :xv i i ) .

D ef in i t ions T he b i o cu l tu ra l a pp roa ch t o h uman s k e le ta l r ema in s v i ew s t h em a s r emnan t s o f as y s t em w h ich w a s i n vo lved r e c ip ro ca l ly w i th i t s n a tu ra l a nd s o c ia l e nv i ronmen t . A t temp t s t oe va lua t e t h e i n t e ra c t ion o f t h e se s y s tem s a r e f r equen t ly m ad e t h rough a na ly se so ft h e h ea l th o fp a s tp opu la t ions , b ecau se h ea l th s t a tu s i nf lu ence s f a c to r s s u ch a s l i f e e xpec tancy , r e p roduc t ive p o ten t ia l a nd t h e c apa c i ty f o r w o rk a nd l e a rn ing , a l o f w h ich a r e c r uc ia lt ot h em a in tenance o f as o c ie ty . F rom e a r l ie s t t i me s , u n t i l t h e i n t roduc t ion o f t h e " g e rm " t h eo ry o f d i sea se b y P a s teu r a nd o t he r s i nt h e l a t e n i ne t een th c en tu ry , d i sea se w a s v i ewed a s ad i s tu rban ce o ft h eb a lance b e tween a n i n d iv idua l a nd h i s e nv i ronmen t , b o th i n te rna l a nd e x te rna l ( Dubo s , 1 9 59 ) . H ippo c ra te s , f o r e xamp le , d e s c r ibed " t he i nf luence o f c l ima te , w a te r s upp ly a nd s i tua t ion o n h ea l th " ( Chadw i ck a nd Mann , 1 9 50 :90 ) . W i th t h e r e cogn i t ion o ft h e l i nk b e tween d i sea se a nd b a c te r ia , t h e l a t te r w e r e c on s ide red t o b e t h e s o le c au se o f t h e f o rme r ( Ca s se l e t a l , 1 960 ; Wy l ie , 1 970 ) . S o , f o r e xamp le , t u be r cu lo s i s w a s s e en a s ac on sequence o fe xpo su re t ot h e t u be r c le b a c i l lu s ,t h es ymp toms o f t h e d i sea se b e ing c au sed b y d i s rup t ion t ot h e p hy s io log ica l s y s tem ; p e r tu rba t ion s o f t h e p sy cho log ica l o r s o c ia l s y s tem s w e r e n o t e n te r ta ined a s p o s s ib le c au s e s o f t h e d i sea se ( Ca s se l e ta l , 1 960 ) . I n r e cen t y ea r s t h e r e h a s

" Hea l th i s t h e p e r f e c t , c on t inu ing a d ju s tmen t o f a n o rgan i sm t o i t s e nv i ronmen t " ( Wy l ie , 1 970 :103 ) . T he d e s c r ip t ion o f h ea l th a s " c on t inu ing " r e in fo r ce s t h en o t ion t h a t t h e c ond i t ion i so ne e nd o f as p e c t rum w h ich a l so i n co rpo ra te s d i sea se . H ea l th " doe s n o t d i sapp ea r d ur ing a n i l ne s s t o r e tu rn o n r e cove ry b u t c on t inue s , e ven t h ough i t m ay d rop i nl e ve l w h i le t h e o rgan i sm i sa dap t ing t o t h e c u r r en t i n su l t " ( Audy , 1 971 :142 ) . A cco rd ing t o Wy l ie ( 1 970 ) , t h e p r inc ipa l s ho r t com ing o f t h e c on t ra s t ing d e f in i t ion o f h ea l th , o f f e r ed p e rhap s m o s t f amou s ly b y t h e Wo r ld H ea l th O rgan i sa t ion , i si t s i n f e r ence t h a t h ea l th a nd d i sea se a r e d i s c r e te a nd m u tua l ly e xc lu s ive c ond i t ion s : " Hea l th i s as t a te o fc omp le te p hy s ica l , m en ta l a nd s o c ia l w e l l -b e ing a nd n o t m e r e ly t h ea b sence o fd i sea se o ri n f i rm i ty " ( P r eamb le t ot h eC on s t i tu t ion o ft h e Wo r ld H ea l th O rgan i sa t ion , 1 946 , q uo ted b y Wy l ie , 1 970 :10 1 ) .

1

s y s tem ( Cox e ta l , 1 983 ) . U nde rt h e i nf luen ce o f w o rk b yC annon a nd S e lye ,a t ten t ion h a s f o cu sed o n t h e p i tu i ta ry -ad r ena l c o r t i ca l s y s tem a nd t h e s ympa the t ic -ad r ena lm edu l la ry s y s tem ( i b id . ) . T he r e a r e t wo a d rena l g l and s , o ne l y ing o ve r e a ch k i dney ,a nd t h ey e a ch c on ta in t wo e ndo cr ine o r gan s , t h e o u t e r c o r t ex , w h i ch s e c re t e s s t e ro id s , a nd t h e i n ne r m edu l la , w h ich s e c re te s t h e c a t echo lam ine s ( a d rena l ine a nd n o rad r ena l ine ) . T he g l and sa r ew e l l supp l ied w i th n e rve sf r om t h e s ympa the t ic s y s t em . T he a d r ena lc o r tex p roduce s t h r ee g r oup s o f s t e ro id h o rmone s : g l uco co r t ico id s , m ine ra lo co r t ico id s a nd a nd rogen s ; t h e e mpha s i s i n s t re s s r e sea r ch h a s b e en o n t h e g l u co co r t ico id s ( t hey f o rm t h e g r ea te s t p a r t o f c o r t ica l s e c r e t ion ) a l though a l t h r ee g r oup s f u nc t ion i n r e sponse t o s t re s s . E igh t ly f i ve p e r c en t o f g l ucoco r t ico id s e c r e t ion i nh uman s i sh yd roco r t i sone , t h e r ema in ing 1 5% b e ing c h ief ly c o r t i co s te rone ( Cox e t a l , 1 983 ) . G luco co r t ico id s ,w h ich a r ee s sen t ia l t ol i f e ,a f f e c t t h e m e tabo l i sm o f p ro t e ins , f a t s a nd c a rbohyd ra te s , i n p a r t icu la r t h e c onve r s ion o f p r o te in s i n to g l u co se a nd i t s s t o rage a s g l y cogen i n t h e l i ve r ; t h ey a l so a f f e c t t h e g r ow th o f c onnec t ive t i s sue . I n h i ghe r c oncen t ra t ion s , a s o c cu r i nc l in i ca l i n te rven t ion o r a s t h e r e su l t o f s t r e s s , g l uco co r t ico id s i n h ib i t p ro te in s y n the s is , b l o ck t h e i n f l amma to ry r e spon se , e l eva t e b l ood s u ga r l e ve l s a nd i n i t ia te t h e l o s s o f c a l c ium a nd p ho spha t ef r om t h ek i dney s( Cox , 1 978 ) . T he a d rena l c o r tex i sr e gu la ted l a rge ly b y t h e b l ood l e ve l o f a d r eno co r t ico t roph ic h o rmone ( ACTH ) , w h i ch i s r e lea sed b y t h e a n te r io r p i tu i ta ry , ag l and s i tua t ed a tt h eb a se o ft h es ku l l . I n r e cen t y ea r s i t h a s b een r e cogn ized t h a t h o rmon e s e c r e t ion i n t h e a n te r io r p i tu i t a ry i s c on t ro l l ed b y t h e h ypo tha lamu s ( a n a r ea o f g r ey m a t te r i n t h e b r a in ) , w h ich s e c re te s c hem ica l a g en t s ; t h e se a r e k nown a s r e lea s ing a nd i n h ib i t ing f a c to r s , b u t a l so a s h ypophy seo t rop ic h o rmone s ( Ganong , 1 985 ) . T he a n t e r io rp i tu i ta ry r e lea se s g r ow th h o rmone a nd p ro la c t in ( am i lk p r oduc ing h o rmone ) a nd f o u r o t he r h o rmone s i n c lud ing ACTH , w h ich c on t ro l t h e a c t iv i t ies o f o t he rg l and s . T hey h ave a l b een a s so c ia t ed w i th t h e s t r e s s r e spon se , a nd m ay p l ay a r o le i n r e s i s tance , a l though " i t i st h e a n te r io r p i tu i ta ry a d r eno co r t ica ls y s tem w h ich a ppea r st od om ina te , i fn o t i n r e a l i ty , t h en c e r ta in ly i n t e rms o f t h e a ten t ion p a id t oi ti nr e sea r ch " ( Cox , 1 978 :60 ) . T he a d r ena l m edu l la , w h i ch i sc l o se ly c onnec ted

Wy l ie d e sc r ibed t h i s d e f in i t ion a s " a symp to t ic " , b ecau se i tv i ew s h ea l th a s ag oa l w h i ch u l t ima te ly c anno tb ea ch ieved . H ow d o w e p rog r e s s f r om e co log ica l d e f in i t ion s o f h ea l th t o t h e r e cogn i t ion a nd e va lua t ion o ft h ec ond i t ion i nt h es k e le ta lr ema in s o f e x t inc t p opu la t ion s? T he v eh ic le i s t h e p hy s ica l t r a ce s o ft h e h uman o r gan i sm ' s a dap t ive r e spon se , w h ich a r e g ene ra l ly d e sc r ibed a s i n d i ca to r s o f s t re s s , o r , l e s s f r equen t ly b u t p r e f e rab ly ,m a rke r so fi n su l t . T h i s i n t roduce s af u ndamen ta lq ue s t ion : w ha t i s s t r e s s ? A s u rvey o f t h e a n th ropo log i ca l l i t e ra tu re r e vea led a w id e a nd s ome t ime s u ndef ined u s e o f t h e t e rm i n r e f e r ence t op a s t p opu la t ion s( Tab le 2 . 1 ) ,w h i ch i ss i gn if ican ts i nce t h e c oncep t o f s t re s s w h ich h a s b een t h e m a jo r i nf luence i na n th ropo log ica lu s age ,n ame ly t h a to f S e lye ( 1 950 ; 1 9 57 ; 1 9 73 ) , i n vo lved a c l ea r d e f in i t ion a s w i l l b e s e en s ho r t ly . T he c l in ica l l i t e ra tu r e r e vea l s t h a tp rob lem s i nd e f in ing s t re s s a r e n o t , h oweve r , p ecu l ia r t oa n th ropo logy . T he p opu la r u s e o f " s t r e s s " i si nt h e s e n se o fn e rvou s o r e mo t iona l s t ra in , a nd t h e r e a r e c u r r en t ly n ume rou s b ook s a nd m agaz ine a r t ic le s d ed ica t ed t oa dv i s ing b u s ine ss e xecu t ive s h ow t oa vo id i t s d e le te r iou s c on sequence s . Ma son ( 1 975 ) n o ted t h a t i n t h e e a r ly y ea r s o f t h i s c en tu ry c l in ica l r e sea r che r s , n o tab ly C annon ( 1 9 14 ; 1 928 ; 1 929 ) , a l so r e ga rded s t re s s i n t h i s l i gh t . A l though C annon c on s ide r ed s t re s s i n ap hy s io log i ca l c on t ex t a s w e l l , p a r t i cu la r ly w i th r e ga rd t o i n te rna lh omeo s ta s i s ,i ti st h e w o rk o fH an s S e lye w h i ch d om ina ted t h i s f i e ld f o r t wo d e cade s a f t e r C annon u n t i lt h ee a r ly 1 9 50 s ,a nd w h i ch h a sb een a dop ted b ya n th ropo log i s t st oi d en t i fy e p i sode so f m e tabo l i ci mba lance . S e lye ' sp i onee r ing w o rk l e d t ot h e f o rmu la t ion o f ac oncep t o fs t r e s s , a nd t h e r e ve la t ion o f i t s p hy s io log i ca l m echan i sms ( Cox e ta l , 1 983 ) . H i st h eo ry o fs t re s si sn o tab le f o ri t s t h r ee b a s ic p r em i se s : t h a t s t r e s s i s ar e sponse t o s t imu l i , t h a t t h er e spon se i sn on -spec if i c , a nd t h a t t h e m an i f e s ta t ion o f t h e r e spon se f o rm s a r e cogn i sab le s ynd rome , t h e G ene ra l A dap ta t ion S ynd rome ( G .A .S . ) . A l though i n am a jo r w o rk p ub l i shed i n 1 946 , S e lye u s ed t h e t e rm " s t r e s s " " i n t h e c onven t iona l s e nse o f s t imu l i , e vo ca t ive a gen t s , o ro u t s ide f o r ce s a c t ing o n t h eo r gan i sm " ( Ma son , 1 975 :9 ) , h i s l a te r w o rk i sc ha r a c te r i sed b y h i s p e r cep t ion o f i ta s ar e spon se t os t imu l i . T he p u rpo se o ft h er e spon se i st h em a in t enance o f h omeo s ta s i s , o r b a lan ce , o f t h e s y s tem . A t t h i s p o in t i t w i l l b e h e lp fu l t o c on s ide r t h e m echan i sm sb yw h ich t h i si sa ch ieved .

c on s id e r ed t o b e i n vo lun ta ry a l though t h i s i d ea h a s b e en c ha l l enged ( Boddy , 1 983 ) . I t i s d i v ided i n to s ympa the t ic a nd p a ra sympa the t i cs y s t em s .

P hys io log ica l C ons idera t ions I ta ppea r s t h a t t h e p hy s io log i ca l r e spon se o f t h e b ody i sp r inc ipa l ly t h a t o fa c t iv i ty w i th in t h e a u tonom ic n e rvou s s y s tem 1 a nd t h e e ndo c r ine 2 1

2

T he a u tonom ic n e rvou s s y tem a c t iva t e s g l and s a nd s moo th m u s c le , a nd i sg ene ra l ly

1 2

T he e ndo cr ine s y s tem c on s i s t s o f t h e d u c t l e s s g l and s w h ich s e c re te h o rmone s . E xamp le s a r e t h e p i tu i ta ry g l and a nd t h e t h y ro id g l and .

p sycho log ica l p ro ce s se s " . N o t e t h a t i ti sn o t t h e r e spon se i t s e l f w h ich i sc on t rove r s ia l , t h e i s sue b e ing i st h e r e sponse i t se lf s t r e s s , o ri si te voked b y s t r e s s ? T he k ey s tone o f S e lye ' s mode l i st h e n o t ion t h a t t h e s t r e s s r e sponse i sn on spec if ic , t h a t i s , i t c an b e p roduced b y aw ide r a nge o f s t imu l i , o r s t r e s so r s :

t o t h e s ympa the t i c n e rvou s s y s t em , p roduce s a d rena l ine a nd n o rad r ena l ine . I n h uman s , a bou t t e n t ime s mo r e a d r ena l ine t h an n o rad rena l ine i s s e cr e ted b y t h e m edu l la ,b u t l e ve l s o ft h e l a t te ri n p l a sma a nd u r ine a r e g r ea te r t h an t h e f o rme r b ecause o f t h e a dd i t iona l s e c r e t ion o f n o rad rena l ine b y t h e s ympa the t ic n e r ve e nd ing s ( Cox e t a l , 1 983 ) . A d rena l ine i n c r ea se s m e tabo l i sm , c au se s t h e l i ve r t or e lea se g l ycogen a s g l uco se , a nd d i la te s t h e s ma l l a r te r ie s o f t h e h ea r t a nd m us c le s , a nd t h e b ron ch ia l t u be s . T he a moun t o f a d rena l ine s e c r e ted v a r ie s , a nd i s e l eva ted b y f e a r , a nge r o r e xc i t emen t . C onsequen t ly , t h e a d r ena l g l and s a r e o f ten r e f e r red t o a s t h e " g land s o f f l i gh t a nd f i gh t " ( A rms t rong a nd J a ck son , 1 972 :265 ) . I nc l in ica l s i tua t ion s ,t h ea c t iv i ty o ft h en eu ro e ndoc r ine s y s tem c an b e m ea su r ed d i r e c t ly , b y a s se ssmen to ft h el e ve lo fh o rmona le xc r e t ion ,f o r e xamp le ( Cox , 1 978 ; S c r im shaw e t a l , 1 966 ) . T he a n th ropo log i s t h a s t o r e ly o n t h e i n d i r e c t e v idence o f n eu ro -endo c r ine a c t iv i ty w h i ch c an b e d e tec t ed i n t h e t e e th a nd b one ; s u ch a c t iv i ty g ene ra l ly c au se s t h e r e lea se o f c a tabo l ic h o rmone s ( t ho se w h ich r e lea se e ne rgy ) , s o t h a t g r ow th , a na nabo l ic p ro ce s s( t ha ti s ,o ne r e qu i r ing e ne rgy ) i si n h ib i ted ( Goodman e ta l , 1 984 ) . B one g r ow th i s ag ood i l u s t ra t ion o ft h i sp roce s s ; b one i s" ah i gh ly s e n s i t ive t i s sue ,a dap t ing i t se l fr a p id ly t o v a r ia t ion s i n t h e g ene r a l w e l l -be ing a nd s o c on s t i tu ted t h a t i n t i me s o f s p ec ia l s t ra in i tc an r e l inqu ish i t s g row th a c t iv i ty e n t i r e ly a nd r e l ieve t h eg ene ra l e conomy b ye l im ina t ion o ft h a ta dded b u rden " ( Pa rk , 1 954 :281 ) . O ne f r equen t ly r e co rded m a rke r o f " s t r e s s " i s d en ta l e name l h ypop la s ia . T h i s o ccu r s w hen t h e s yn the s i s o f p ro t e in , t h e b a s ic b u i ld ing -b lo ck o f t h e b ody , i s d i s rup ted b y e l eva ted l e v e l s o f h yd ro co r t i sone f o l low ing a c t iv i ty o f t h e a d r ena l c o r tex . T h i s i n t u rn c ause s t h e r e duc t ion o r c e s sa t ion o f f o rma t ion o f t h e e name l p ro te in m a t r ix b y a me lob la s t s ,p roduc ing ah ypop la s t ic l e s ion ( Ro se e ta l , 1 985 ) .

" By n on sp ec if ic w e m ean e f fe c t s o r r e spon se s t h a t a r e c ha ra c te r i s t ic o f m any s t imu l i o r a gen t s -f o r e xamp le , t h e m an if e s ta t ion s o f t h e a l a rm r e a c t ion , w i th t h e s e c r e t ion o f a d r eno co r t i co t roph ic h o rmone ( ACTH ) , c o r t ico id s a nd c a t echo lam ine s . . . " ( Se lye , 1 976 :53 ) . I n h i s f i r s t p ub l ica t ion S e lye d e sc r ibed s t r e s so r s a s " no cuou s " ( 1 936 :32 ) , b u t l a t e r r e po r ted t h a t p l ea su rab le s t imu l i " . .a g ame o f t e nn i so re ven ap a s s iona te k i s s . . . " ( 1957 :53 ) m ay e l i c i t t h e s t r e s s r e sponse . S ub sequen t s t ud ie s h ave a l so s hown t h a t e xpe r ience s o f ag ene ra l ly e n joyab le k i nd a r e a lmo s t a s e f f ec t ive a s u np lea san t e xpe r ience s i n i n ten s ify ing c a te cho lam ine ( e .g . a d rena l ine ) e xc r e t ion ( Cox , 1 9 78 ) , b u t t h i s f i nd ing i sr a r e ly r e f e r red t oi nt h e c l in ica l l i t e ra tu r e a nd , a s f a r a s Ia m a wa r e , h a s n o tb een n o ted i na n th ropo log i ca ls t ud ie s . T he n o t ion o f n on spe c i f ic i ty h a s b een q ue s t ioned . F o r e xamp le , i ti sp ropo sed t h a t t h e g ene ra l n on spe c if ic p a t t e rn o f n eu roendo c r ine r e spon se t os t r e s so r sc anno tb ed emon s t r a ted w i th t h e f a c i l i ty w h ich S e lye p ropo sed , s p ec if ic p a t ter n s o f r e ac t ion h av ing b een s hown t o b e p r oduced b y d i f f e r en t s t re s so r s ( Cox e ta l , 1 983 ) . N ume rou sd i f f e r en tt y pe so fp hy s i ca l s t imu l ih ave b een s hown t oi n c r ea se t h ea c t iv i ty o ft h em edu l la ( Cox e ta l , 1 983 ) ,b u ta ga in s t t h ec oncep t o fn on s p e c i f ic c a techo lam ine s e c r e t ion i st h e h ypo the s is t h a t a nx ie ty -p roduc ing s i tua t ion s a r e c onnec ted w i th a d r ena l ine r e lea se , w h i le n o rad r ena l ine i s s e c r e ted i n c i r cums tance s o f a gg r e s s ion ( Cox , 1 9 78 ) . I n i t ia l ly , S e lye f o cu sed o n p hy s i ca l s t r e s so r s ( s uch a s i n f ec t ion , i n ju ry , c o ld o r h ea t ) t ot h e e x ten t t h a t C ox ( 1 987 ) c l a imed h e i g no red t h e p a r t p l ayed b y p sycho log ica l p roce s se s , a l though i n al a te r p ape r S e lye s t a ted t h a t i n h uman s " p sycho log ic s t re s so r s , a nd p a r t icu la r ly e mo t iona l a r ou sa l , a r e o f p r ima ry i mpo r tance . T h i s i s e spe c ia l ly t r ue o f t h e mo s t c ommon s t r e s so r s o fd a i ly l i f e ( f ru s t ra t ion , a nge r , f e a r a nd h a t e ) " ( 1 976 :56 ) . Ma son ' s ( 1 974 ) e xpe r imen ta l w o rk w i th monkey s i n d ica ted t h a t t h e p i tu i ta ry a d r ena l c o r t i ca l a x i s d o e s n o t r e a c t n on s p e c i f ica l ly t op hy s ica l s t imu l i s u ch a s h ea t a nd f a s t ing , a nd t h a t i n t h e c a se o f f a s t ing , f o r e xamp le , i ti sa ppa r en t ly t h e s i tua t ion a s aw ho le , w i th i t s u np lea san t p sycho log ica l a spe c t s , w h i ch e l ic i t s ar e spon se , r a the r t h an t h e s imp le a b sence o ff o od . T hu s Ma son p ropo sed t h a t t h e n o t ion o f p hy s io log ica l " non -spe c i f ic i ty " w h i ch i s f u nd amen ta l t oS e lye 's t h eo ry i su nde rm ined , f o r i t m ay b e t h a t t h e n on sp ec i f ic r e sponse i s

S e lye 's Mode l o fS tre s s We c an n ow l o ok i n al i t t le m o r e d e ta i l a t S e ly e ' s mode l o fs t r e s s , a nd e xam ine s ome o ft h e c r i t i c i sms w h ich h ave b een l e ve l led a t i t . T o r e cap i tu la te , S e lye p ropo sed s t r e s s t o b e an on s p ec i f i c r e spon se t os t imu l i , t h e m an i f e s ta t ion o f t h i s r e sponse f o rm ing t h e G ene ra l A dap ta t ion S ynd rome ( F igu re 2 . 1 ) . H i s n o t ion o fs t re s s a sa r e spon se , r a the r t h an a s as t imu lu s , i sn o t s ha r ed b y m o re c on t empo ra ry r e sea r che r s . S i gn if ican t ly f o r t h e b i ocu l tu r a l a pp roa ch , C ox a nd h i s c o l league s ( 1983 :256 ) s umma r i sed c r i t i ca l v i ew s o fs t r e s s a s ar e spon se b y d e s c r ib ing s u ch m ode l s a s d e f ic ien t w hen c ompa r ed t o c u r r en t " p sycho log ica l a nd p sy chophy s io log ica l mode l s o f Man " w h ich e mpha s i se t h e" dynam ic n a tu re o f t h e p e r son ' s i n te rac t ion w i th h i s e nv i ronmen t , a nd . . . the m ed ia t ing r o le o f i n d iv idua l

1 3

T he G .A .S . h a s t h re e d i s t inc t p ha se s ( F ig .1 ) . T he f i r s t ,t h ea l a rm r e a c t ion ,i sc h ief ly a ni n c r ea se i na c t iv i ty o f t h e s ympa the t i c -ad rena l m edu l la ry a x i s , a nd i sa c compan ied b y ar e duced l e ve l o f r e s i s tance , w h ich m ay c o l lap se i ft h e s t r e s so r i s s u f f i c ien t ly s e ve r e , i n w h ich c a se d ea th w i l l e n sue . H oweve r , i fa dap ta t ion i sp o s s ib le i nt h e f a ce o f c on t inued e xpo su r e t o t h e s t r e s so r , t h e s e cond p ha se , t h a t o fr e s i s tance , w i l l b e e n t e red . T he b od i ly c hange so ft h e f i r s ts t age a r er e p laced b y t h o se i n d ica t ive o fa dap ta t ion , s uch a s ar e tu rn t on o rma lb ody w e igh t( S e lye , 1 973 ) ; t h i s s e cond p ha se i s p r ima r i ly o ne o f i n cr ea sed a n te r io r p i tu i ta ry -ad r ena l c o r t i ca l a c t iv i ty ( Cox e t a l , 1 9 83 ) . L ong t e rm e xpo su r e t os t re s s u s e s u p t h e e ne rgy n eeded f o r a dap ta t ion t o i t , s o t h a t t h e t h i rd p ha se ,t h a to fe xhau s t ion ,w i l lo ccu r ,l e ad ing t od ea th . O n ly t h em o s t s e ve r es t re ss l e ad st ot h e s t age o f e xhau s t ion ; m o s t s t re sso r s p r oduce c hange s w h i ch c o r r e spond t ot h e f i r s t t wo s t age s o n ly , af r equen t ly e xp e r ienced s i tua t ion i nn o rma l h uman l i f e ( S e lye , 1 957 ) . S t re s s e x i s t s a ta ny g i ven t i me d u r ing t h e t h r ee s t age s o f t h e G .A .S . S e lye ( 1 957 :54 ) e mpha s i sed t h a t s t r e s s a nd t h e G .A .S . a r e n o t s y nonymou s , b e cau se " t he se r e a c t ion s a r e c h a ra c te r ized b y c e r ta in m ea su rab le o r gan -change s w h ich a r e c au sed b y s t r e s s a nd h ence c ou ld n o tt h em se lve sb es t re s s " . A cco rd ing t oC ox a nd h i s c o l league s ( 1 983 ) , t h e G .A .S . d oe s n o td e sc r ibe p r ec i se ly t h eb ody ' s r e spon se t os t r e s s . R e f e r r ing t o Ma son ' s ( 1 968 ) w o rk , t h ey p r opo sed t h a t w ha t h app en s i nt i me s o fs t r e s si si n c r ea sed p a t t e rn so fc a tabo l ic a c t iv i ty , a nd t h e r e ve r sa l o f t h e s i tua t ion w hen s t r e s s i s a me l io ra t ed . Ma son ( 1 971 :330 ) a r gued t h a t t h e G .A .S . i si n compa t ib le w i th t h e w e l l -e s tab l i shed p r inc ip le so fh omeo s ta t ic r e gu la t ion : " h ow c ou ld t h e s ame h o rmona lr e spon se h ave a dap t ive u t i l i ty i n r e spon se t o d i ve r se s t imu l i w h i ch p l ace d i ame t r ica l ly o ppo s i te m e tabo l ic n eed s o n t h e b ody a sd o , f o re xamp le , h ea t a nd c o ld ? " , a nd h e b e l ieved t h e e xp lana t ion t ob e ag ene ra l a dap t ive r e spon se o f b ehav iou ra l o rp sycho log ica l o r ig in . A f i na l b u t i mpo r tan t p o in t t o n o te i st h e p a r t p l ayed b y t h e h ypo tha lamu s i nt h e s t imu la t ion o f h o rmona l s e c r e t ion ; f o re xamp le , af a i lu r ea tt h i s l e ve l w i l l c a u se f a i lu r e o fh o rmona l s e c r e t ion b y t h e a n t e r io r p i tu i ta ry ( Rus s e l l , p e r sona l c ommun i ca t ion ) . T h i s i ss i gn i f ican t i nt e rms o f t h e l e ve l a t w h i ch S e lye i d en t if ied t h e b ody ' s r e spon se t os t r e s so r s . T he e f f e c t o fS e lye ' s c on t rove r s ia l t h eo ry h a s b een e ncap su la t ed b yE nge l :

b ehav iou ra l , r a the r t h an e ndo cr ine . T he p sy cho log ica l c omponen t r a i se s a f u r the r c r i t ic i sm o fS e lye ' s h ypo the s i so fn on spec i f i c i ty . S e lye h im se lf r e cogn i sed t h a t h i s t h eo ry w a s w eakened b y t h e f a i lu r e t o r e cogn i se t h e m echan i sm b y w h ich s o m any d i f f e r en t s t re s so r s c ou ld p a s st h em e s sage t on eu roendo c r ine c en t re s o f t h e n eed f o r a n a dap t ive r e spon se ( Ma son , 1 971 ) . S e lye ( 1 976 :54 ) c a l led t h i s u nknown m echan i sm t h e " f i r s t m ed ia to r " o f s t r e s s r e spon se s w h ich Ma son ( 1 971 ) h a s s u gge s ted m ay b e j u s t t h e p sycho log i ca l e qu ipmen t c once rned w i th s t imu la t ion o re mo t iona lr e ac t ion s t o d ange rou s o r d i sag re eab le c i r cums tance s i n e ve ryday l i f e . T h i s i s n o t t o s a y t h a t p sycho log i ca lp roce s se sa r en o tp hy s io log ica l ,b u t r a the r i th i gh l igh t s t h e l e ve l w i th in t h e c en t r a l n e rvou s s y s tem a t w h ich i n t eg ra t ion o f n eu ro endo c r ine r e spon se s o c cu r ; t h e l owe r l e ve l e ndoc r ine r e spon se s t o h ea t o r e xe r c i se , f o r e xamp le , a r e i n g ene ra l m o r e c on s i s ten t i n te r i n d iv idua l ly t h an a r et h eh i ghe rl e ve lr e spon se st o d i s t inc t " p sycho so c ia l s t imu l i o r l i f e e ven t s " ( Ma son , 1 975 :32 ) . S e lye ( 1 975 ) c oun t e r ed Ma son ' s i n te rp r e ta t ion o f t h e " f i r s t m ed ia to r " a s p sy cho log i ca l o n t h eg r ound s t h a ts t r e s so c cu r si n l owe r a n ima l s a nd p l an t s l a ck ing a n e rvou s s y s t em , a l though h e o f f e r ed n o o pe ra t ive m echan i sm ; i n a dd i t ion , h e p o in ted t o s t r e s s p r oduced b y a nae s the s ia , w h ich h e p r opo sed i sa w e l l -known s u rg ica lp henomenon . I tn ow a ppea r s t h a t t h e m a jo r i ty o fh o rmone s , p o s s ib ly a l o ft h em , c ha ra c te r i s t ica l ly r e spond t o m u l t ip le s t imu l i ( Ma son , 1 975 :27 ) , a nd i n t h i s s e n se t h e r e sponse o f h o rmone s s u ch a s a d r ena l ine , g r ow th h o rmone a nd h yd ro -co r t i sone c ou ld b e s a id t o b e " non sp ec if i c " i n t h a t t h e s e c re t ion o fe a ch m ay b es t imu la t ed b ym o r et h an o ne s t imu lu s . H oweve r , Ma son h a s f o und n o e v id ence " t ha ta ny s i ng le h o rmone r e spond st oa l l s t imu l i i n a bso lu te ly n on spec if i c f a sh ion , a s i mp l ied i nS e lye ' sd e f in i t ion o fs t re s sa so ccu r r ing i nr e spon se ' t oa ny d emand- . T he t h i rd b a s ic p r em ise o f S e lye ' s s t r e s s t h eo ry i s t h e G ene ra l A dap ta t ion S ynd rome ( G .A .S . ) , w h ich p r opo se s t h a t a l though t h e s t r e s s r e spon se m ay b e t r i gge r ed b y aw ide r a nge o f s t r e s so r s , a nd i s t h u s n on spe c i f ic , t h e m an if e s ta t ion o f t h e r e spon se i sh i gh ly s p e c i f ic a nd f o rm s ar e cogn i sab le s y nd rome : " S t re s s i s t h e s t a t e m an i f e s ted b y a s p e c i f ic s ynd rome w h ich c on s i s t so fa l t h e n on spec if i ca l ly i n duced c hange s w i th in a b i o log ic s y s tem . T hu s s t r e s s h a s i t s o wn c ha ra c te r i s t ic f o rm a nd c ompo s i t ion b u tn o p a r t icu la rc au se " ( S e lye , 1 9 57 :54 ) .

" I n h i s t h eo ry o f s t re s s . .P ro f e s so r S e lye h a s p r e sen t ed a c oncep t w h ich i s a lmo s t b r ea th tak ing i ni t s s cope . I th a s e l i c i t ed e x t r avagan t p r a i se a nd c r i t ica l c ondemna t ion . I th a s p e rmea t ed m ed i ca l t h ink ing a nd i n f luenced m ed ica l r e sea r ch i ne ve ry l a nd , p r obab ly mo re r a p id ly a nd m o re i n t en se ly t h an a ny o t he r t h eo ry o f

S e lye ( i b id . ) d e s cr ibed t h i s a sa n " op e ra t iona l d e f in i t ion ; i tt e l l s w ha t m u s t b e d one t op r odu ce a nd r e cogn ize s t re s s " ;i ti st h em an i f e s ta t ion so fa s t a te w h i ch m ak e i tr e cogn izab le , s o i tf o l low s t h a tt h em an if e s ta t ion so ft h es t re s ss y nd rome ,t h e G .A .S . ,a low t h es t a te o fs t re s st ob ei d en t i f ied .

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d i sea se e ve rp ropo sed " ( Enge l , 1 956 , c i ted b y Ma son , 1 975 :10 ) .

p r ov ide af u r the r e xamp le o f t h i s v i ew o f s t re s s ; w hen s t anda rd s o f g r ow th d e r ived f r om l i v ing p opu la t ion s " a r e u s ed i n c on junc t ion w i th a n th ropome t r ic a nd r o en tgenog r aph ic d a ta o n t h e g r ow th r e spon se o f c h i ld r en s u f f e r ing f r om m a lnu t r i t ion , d i sea se a nd o t her p hy s io log ica l s t re s se s , af u l le r u nde r s tand ing o f g row th a nd m o rb id i ty a mong p r eh i s to r i c p eop le s i s f a c i l i ta ted . S uch i n fo rma t ion m ay p rov ide i mpo r tan t i n s igh t s i n to t h e b i o cu l tu ra l a dap ta t ion o fe a r l ie rh uman s o c ie t ie s " ( my e mpha s i s ) . I st h i s a n a dequa te v i ew o f s t r e s s u pon w h ich t o b u i ld b i o cu l tu ra l i n te rp r e ta t ion so fp a s t p opu la t ions ? A r e v iew o ft h e c l in i ca l l i t e ra tu r e i n d ica tes t h a t i ti s n o t .

S e lye w a s l a rge ly r e spon s ib le f o r t h e s u rge o f i n te re s t i n p hy s io log ica l s t r e s s i n t h e 1 950 s y e t n ow , al i t t le m o re t h an t h i r ty y ea r s a f te r E nge l ' s s t a temen t , h i s w o rk i sl a rge ly o ve r shadowed b y s ub sequen t r e sea r ch w h ich h a s r e vea led t h e c omp lex i ty o f t h e n eu roendo c r ine r e spon se ( Ru s se l l , p e r sona l c ommun i ca t ion ) , a nd b y a t u rn ing a way f r om p hy s io log ica l t op sycho so c ia l s t r es sr e sea r ch ( Ma son , 1 9 75a ) . T he d eve lopmen t i nt h e e a r ly 1 950 s o f m o re s e n s i t ive a nd p r e c i se m e thodo log ie s t h an h ad b een a va i lab le t o S e lye c onf i rmed h i s f i nd ing s o f e l eva t ed l e ve l s o f c o r t ico s te ro id s i nb l ood a nd p l a sma i nr e spon se t o av a r ie ty o fs t imu l i . A s ar e su l t , a l though i tw a s l a te rc en su r ed , t h e r e w a s aw ide a c cep tance o ft h e c oncep t o f a n on spe c if ic r e spon se b y t h e p i tu i ta ry -ad r ena l c o r t ica l s y s tem , a nd ad e c l ine i n t h e t e s t ing o f h ypo the se s o f p hy s io log ica l s t r e s s ( Ma son , 1 971) .

P sycho log ica l I nf luence so n H ea l th T he r e la t ion sh ip b e twe en h ea l th a nd a n a dve r se p hy s ica l e nv i ronmen t i sw e l l -known , a nd d emon s t ra ted ,f o ri n s tance , b yt h ef a l l i nt h ed ea th r a te c on sequen t u pon t h e i n t roduc t ion t o V ic to r ian L ondon o f a dequa te s ewe rage a nd a c l ean w a te r s upp ly ( Po r te r , 1 987 ) . I th a s b een a r gued c onv inc ing ly ,b y D ubo s( 1 96 1 )a nd H ink le ( 1 96 1) f o r e xamp le , t h a t t h e s o c ia l e nv i ronmen t a l so h a s f a r r ea ch ing c on sequence s f o r h ea l th : " o ne f i nd s n o i l ne s s w h ich i sn o t i ns ome d eg r ee i n f luenced b yt h ew ay t h a t m en r e a c tt ow ha tg oe s o n a r ound t h em , a nd n one t h a t d oe s n o t o ccu r s ome t ime s i na s so c ia t ion w i th m an i f e s ta t ions o f mood , t h ough t o r b ehav iou r ( H ink le , i b id :295 ) . I n he r en t i n s u ch r e a c t ion s a r e p a t te rns o f a dap ta t ion w h ich a r e r e gu la ted b y t h e a c t iv i ty o f t h ee ndo c r ine g l and s , t h e n e rvou ss y s tem a nd " t he g r o s sb ehav iou ro ft h ep e r son " ( i b id . :294 ) . I n d iv idua l s i n te rac t w i th t h e i r s o c ia l e nv i ronmen t a td i f f e r en t l e ve l s : a ta n i n t ima te , i n te r -pe r sona l l e ve l a nd m o re g ene ra l ly w i th t h e s o c ie ty o r c ommun i ty t o w h ich t h ey b e long . P sycho log ica l s t r e s s m ay b e e xpe r ienced a tb o th l e ve l s . G i lbe r t( 1 985 ) p ropo sed t h a ta l though i ti s a lway s p r e sen t i n s o c ia l c ommun i t ie s , i ti sl e s s p r eva len t i n s t ab le s o c ie t ie s t h an i n t h o se e xpe r ienc ing s o c io cu l tu ra l c hange . H e i l u s t r a ted h i s i d ea b y r e f e r ence t o Mo s s ' ( 1 973 ) mode l o f s t r e s sa s ar e su l to ff a i lu r et oc ommun ica te :

A n thropo log ica l C oncep ts o f" S tres s" T he d e c l ine i n c l in ica l i n te r e s t i n p hy s io log i ca l s t re s s h a s b een m a t ched b y a n e n thus ia s t ic u p take o n t h ep a r t o fa n th ropo log i s t s . B u ik s t ra a nd C ook ( 1 980 ) s aw t h i s a s ar e spon se t oc r i t ic ism so ft h ev a lue o fs k e le ta ls t ud ie sb y t h e " n ew " a n th ropo log i s ts . S o o nce a ga in S e lye h a s b een i n f luen t ia l , f o r h i s i st h e c oncep t o f s t r e s s w h ich f o rm s t h e b a s is , e xp l ic i t ly o ri mp l ic i t ly , o f m any i n te rp r e ta t ion s o f s t r e s s i nt h e s k e le ta l a nd d en ta l r ema in s o f p a s t p opu la t ion s ( e .g . B u ik s t r a a nd C ook , 1 980 ; C la rk , 1 985 ; C la rke , 1 978 ; C ondon , 1 981 ; G i lbe r t , 1 985 ; G oodman e ta l , 1 9 80 ; G oodman e ta l , 1 984 ; Ma r t in e ta l , 1 985 ; P owe l l , 1 985 ; R o se a nd P a s ley , 1 980 ; R o se e ta l , 1 9 85 ) . I n t h e i r s u rvey o f Ame r ican p a laeopa tho logy , B u ik s t ra a nd C ook ( 1 980 :444 ) d e s c r ibed t h e i n c r ea s ing u s e o f i n d ica to r s o f s t r e s s , s t a t ing " S t r e s s ' i n t h i s s e n se i s S e lye ' s c oncep t : a ny e x t r in s ic v a r iab le o rc omb ina t ion o f v a r iab les w h ich c au se s t h e o r gan ism t o r e a c t " . Wha t t h ey h ave , i n f a c t , d e s c r ibed i s S e lye ' s n o t ion o f a" s t re s so r " , r a the r t h an s t r e s s , b u t t h e p o in t o f t h i s r e f e r ence t ot h e i r w o rk i st h a t t h ey s e e S e lye ' s m ode l a s c en t ra l t o a n th ropo log ica l i n te rp r e ta t ion so fn on spe c if ic s k e le ta lm a rk e r s . W i th t h e e xcep t ion o fC la rk ( 1 985 ) , t h e r e h a s b een n o p ub l i shed r e cogn i t ion o f t h e c r i t ic i sm s w h ich h ave b een l e ve l led a tS e lye ' s m ode l , a nd t o m yk now ledge , o n ly C la rk ( 1 985 ) , G i lbe r t ( 1 985 ) a nd H i l lson ( t h i s v o lume ) h ave r e f e r r ed e xp l ic i t ly t o p sycho log ica l s t re s s ( d ef ined b y Ma son ( 1 975a :11) a s " t he i mpa c t o f p sycho so c ia l i n f luence s u pon t h e o rgan i sm " ) , a n th ropo log i s t s h ab i tua l ly u s ing " s t r e s s " i n t h e s e n se o f a p hy s io log ica l p henomen ( T ab le 2 . 1 ) : " S t r e s s i sa c e n t ra l , s t e r eo typ ic p hy s io log ica l r e spon se " ( Goodman e ta l , 1 980 :515 , c i t ing L ev i , 1 9 72 , m y e mpha s i s ) . H umme r ta nd V an G e rven ( 1 983 :471 )

" . .be cau se e a ch i n d iv idua l i s e nme shed i n s ome s o c ia l n e two rk ( s ) , a t a ny t i me t h e i n d iv idua l f a i l s t o i n te rp re t c o r re c t ly t h em ean ing a nd a c t ion o fo t he r s , h i s p rob ab i l i ty o f c o r r e c t r e sponse i s r e duced , c on sequen t ly i n c rea s ing h i s d eg re e o f a nx ie ty a bou t r e spond ing a pp rop r ia te ly . T h i sa nx ie ty i st r ans fo rmed i n to a c t iva t ion o f t h e s ympa the t ic n e rvou s s y s t em w i th a t tendan t p roduc t ion o f a d r ena l ine t h a t o ve r t i me r e duce s h i s r e s i s tance t o d i sea se . I fd u r ing t i me s o f c u l tu r e c hange t h e i n d iv idua l p a r t ic ipa te s i nv a r iou s n e two rk s t h a t a r e i n compa t ib le o r f a i l t op rov ide t h e p rope r s o c ia l c l ue s

1 5

f o r b ehav iou r , h i s d eg r ee o f s t re s s w i l l i n c r ea se "( G i lbe r t , 1 985 :34 1) .

s t udy s how st h a t ad i e tm ay b en u t r i t iona l ly s o und b u t c on sumed i n u nf avou r ab le p sycho log ica l c ond i t ion sw h i ch i n h ib i tg r ow th . B e cau se m a rke r s o f i n su l t c an f o rm o n ly w h i le g r ow th i so ngo ing , t h ey r ef l ec t m e tabo l i c i mba lance i n c h i ldhood ; p eak f r equenc ie s o f m a rk e r s i ni n f an t s a nd y oung c h i ld ren h ave b een a t t r ibu t ed t o a" wean ing c r i s i s " ( e .g .C a r l son e ta l , 1 9 74 ; C ook , 1 9 79 ; R o se , 1 977 ; R udney , 1 9 83 ) , b r ough t a bou t b y t h e s yne rg i s t ic i n te ra c t ion o f s u ch f a c to r s a s e x t end ed l a c ta t ion b y u nde rnou r i shed m o the r s , a n u n su i tab l e o r n u t r i t iona l ly i n adequa t e w ean ing d i e t a nd w ean l ing 4 d i a r rhoea . I t may b e t h a t p sy cho log ica l f a c to r s a r e ap a r t o f t h i s " wean ing c r i s i s " a nd c on t r ibu te t ot h e f o rma t ion o fm a rke r s o fi n su l t , e s pe c ia l ly i n ac h i ld a l ready p hy s ica l ly d eb i l i ta t ed . C onf l ic t b e tween p a ren t a nd c h i ld m ay o ccu r a tw ean ing , e i the r b ecau se t h e i n f an t w i she st oc on t inue b r ea s tf e ed ing o ra s ar e su l to f s i b l ing r i va l ry ( Sm i th , p e r sona l c ommun i ca t ion ) . E n fo r ced w ean ing m ay n o t b e t h e o n ly c hange w h i ch r e su l t s f r om t h e b i r th o f as i b l ing ; s t ud ie s h ave s hown t h a t b ehav iou ra l c ha nge s a r e c ommon ,e spe c ia l ly w i th r e ga rd t ot h ei n t e ra c t ion o fm o the r a nd c h i ld ( Dunn a nd K end r i ck , 1 9 82 ) , t h ea r r iva lo f an ew b aby o f ten p r es su r i s ing af i r s t b o rn i n to g r ea t e r i n dependence ( Dunn , 1 9 84 ) . T he f i r s t b o rn i sm o re l i ke ly t h an i t s y ounge r s i b l ing t o b e r e p r imanded w hen a r gumen t s b e twe en s i b l ing s o c cu r , a nd t h e r e i sf r equen t ly a m a rk ed r e duc t ion i nt h e a moun t o fa t t en t ion p a id t o af i r s tbor n w i th t h e b i r th o f as i b l ing ( i b id . ) . A ny o f t h e s e f a c to r s c ou ld h ave a n a dve r se p sy cho log ica li mpa c t ,w h i ch c ou ld d i s rup tg r ow th a nd t h u s i nf l uen ce t h e f o rma t ion o f m a rke r s o f i n su l t . Af i na l , i n t r igu ing p o in t t on o t e i st h a t t h e l i t e ra l t r an s la t ion o f " kwa sh io rko r " ( t he p r o te in d e f ic iency d i sea se c ommon i n d eve lop ing c oun t r ie s )i s" f i r s t se cond " ( FAO /WHO /UN ICEF , 1 8 70 :27 ) ; i t i s t h e d i sea se o f t h e t o dd le r s u pe r seded , w h i le s t i l l b e ing b r ea s t f e d , b y t h e b i r th o fa no the rb aby . I nt r i ba l s o c ie t ie s m o the r s o f ten n u r se t h e i rc h i ld r en f o rt h ree y ea r so rm o r e , s o t h a t t h e c h i ld i sl i v ing i n as t a t e o fi n t ima te d ependen cy o nt h em o the r . Mo the r su s ua l ly w ean w hen t h ey k now t h a tt h ey a r ep r egnan t ,s ot h a tt h e c h i ld m ay a d ju s t , b u t a l though r e la t ive s t a ke o n m uch o f t h e c a r e o f t h e c h i ld a t t h i s t i me , t h e m o the r -ch i ld b ond r ema in s s t rong ; w hen t h e s i b l ing i s b o rn , h oweve r , t h e mo the r ' s w ho le a ten t ion i s t u rned t o t h e n ewbo rn ( E ib l E ibe s f e ld t , 1 9 89 ) . O f c ou r se , p sycho log ica l f a c to r s c a nno t b e e x t ra c t ed f r om a r cha eo log ica ld a ta i nt h ew ay t h a t

S ke le ta l a nd d en ta l m a rke r s r e p r e sen t i n su l t s e xpe r ienced d u r ing t h e y ea r s o f g r ow th , a nd u nf avou rab le f am i l ia l c i r cums tance s m ay h ave a p ro found i mpa c t u pon t h e g r ow ing c h i ld . T h i s i s d emon s t ra ted c l ea r ly b y a s t udy o f t h i r teen c h i ld r en w i th u n s tab le h ome l i ve s w ho p r e sen t ed w i th s ymp tom s s u gge s t ive o f i d iopa th i c h ypop i tu i ta r ism 3,n o tab ly s ho r t s t a tu re ( Powe l l e t ( 2 / , 1 967 ) . Ar e duc t ion i nr e lea se o f ACTH a nd g r ow th h o rmone w e r e t h e m o s t m a rked d e f ic ienc ie s n o ted . R emova l t o a m o r e f a vou r ab le e nv i ronmen t l e d t oa n a c ce le ra t ion o f g r ow th w i thou t t h e a dm in i s t ra t ion o f g r ow th h o rmone . A cco rd ing ly i tw a s p o s i ted t h a t t h e " emo t iona l d i s tu rbance i n t h e se c h i ld r en m ay h ave h ad a n a dve r se e f f ec t u pon r e lea se o f p i tu i ta ry t r op ic h o rmone v i a t h e c en t ra l n e rvou s s y s t em " ( i b id . :1282 ) w h ich w a s m an i f e s t ed i n g r ow th f a i lu re . F u r the r e v idence o f t h e e f f ec t s o f a n u n f avou rab le s o c ia l e nv i ronmen t h a s b e en p r e sen ted b y W iddow son ( 1 951 ) i n h e r f amou s s t udy o f o r phanage c h i ld r en i n p o s t -wa r G e rmany . S he o b se rved r e duced w e igh t g a in , d e sp i te c a lo r i f ic s upp lemen ta t ion , i n c h i ld r en s ub jec ted t o" h a r sh a nd u n sympa the t ic h and l ing " ( i b id . :1318 ) b y t h e h ead o f t h e i r o r phanage ; b y c on t ra s t , e i gh t f a vou r ed c h i ld ren g a ined m o r e t h an d oub le t h es t anda rd a moun to fw e igh to n t h e s upp lemen ted d i e t . I nt h i ss t udy ,g r ow th i nh e igh t w a sn o ta f f ec ted b ye mo t iona l f a c to r st ot h e s ame d eg r ee a sw e igh t ,b u ts im i la rt r end sa r ea ppa r en t . T he i n f luence o f p sy cho log ica l f a c to r s o n p r o te in m e tabo l i sm h a s b een d emon s t ra ted b y S c r im shaw a nd c o l league s ( 1 966 ) . T he i r i n ve s t iga t ion o f 2 6 m a le s t uden t s b e fo r e a nd d u r ing f i na l e xam ina t ion s r e vea led t h a t i n 1 5 c a se s ,d e sp i te c on s i s tency o fd i e t , t h i sp e r iod w a s a s so c ia ted w i th m e tabo l i c c hange s , n o tab ly e l eva t ed n i t rogen e x c r e t ion , i n d ica t ing t h e s e c r e t ion o fc a tabo l ic h o rmone s . A l though t h i s i s as t udy o fa du l t s w ho se g r ow th o bv iou s ly w ou ld n o tb ei nf luen ced ,i ti sa dd i t iona lt e s t imony o ft h e p owe r o fp sycho log i ca l f a c to r s t oa f f e c t p hy s ica l p r o ce s se s . G iven t h a tt h ec au se o ft h ed i s tu rbance sw h i ch l e ave t h e i r t r a ce s i n d en ta l a nd o s seou s t i s sue c anno t b e c a tego r ica l ly i d en t i f ied , h ow i mpo r tan t i si tt o r e cogn i se t h a tp sy cho log i ca l f a c to r s m ay p l ay ap a r t i nt h e f o rma t ion o fm a rke r s o fi n su l t ? Ib e l ieve i tt ob e e s sen t ia l b e cau se m a rke r s h ave b een i n t e rp r e ted i nt e rm so fn u t r i t iona l s t a tu sa nd o fs ub s i s tence c hange , p a r t icu la r ly w i th r e ga rd t o t h e t r an s i t ion f r om h un t ing a nd g a the r e r ing a sa m ode o f s ub s i s t en ce t o o ne o f f o od p roduc t ion ( e .g . C ohen a nd A rme lago s , 1 984 ) ; W iddow son ' s

4

3 Ac ond i t ion r e su l t ing f r om i n su f f i c ien t p i tu i ta ry s e c r e t ion .

1 6

A" wean l ing "i s ac h i ld b e tween t h et i me t h a t af o od o t he rt h an b r ea s tm i lk i s c on s i s t en t ly t a ken a nd t h et i me t h a ti t c omp le te st h et r an s i t ion t o ag ene ra ld i e t ( Go rdon , 1 969 :220 ) .

c hange s i n s ub s i s t ence m ay b e , a l though c i r cum s tance s w h ich c ou ld b e e xpe c ted t oi n du ce p sycho -soc ia l s t re s s i n ap opu la t ion ( b ea r ing i n m ind t h e i n d iv idua l i ty o f r e spon se ) m igh t b e i d en t i f iab le .

s y s t ema t ic a s se s smen t o f t h o se a spec t s o f e xpe r ience t h a t m ake h im u n ique " ( i b id . :292 ) s hou ld a l so b e u nde r taken i si mpo s s ib le , s i nce t h ey a r eu nknowab le t oa r cha eo log i s t s .

C onc lu s ions

P o s tscr ip t : T he I nd iv idua l i ty o f R e sponse

T h i sc hap t e rh a se mpha s i sed t h en eed t oa low t h a t p sycho log ica l e l emen t s h ave t h e p o t en t ia l t o i n f luence t h ef o rma t ion o fm a rke r so fi n su l t . I ti s s u gge s ted t h a t t h e u s e o fS e lye ' s m ode l o fs t re s s h a sl e dt o af a i lu re ,b ya nd l a rge , t oc ons ide rs u ch e l emen t s , t h e c on sequence o f w h i ch i s t h a t m a rke r s o f i n su l t m ay h ave b een i n te rp re t ed e r roneou s ly . T h i si nt u rn h a si mp l i ca t ion s f o rt h e w ide r a pp re c ia t ion o f p a s t s o c ie t ie s . Mo re p r o sa i ca l ly , Iw ou ld a r gue t h a ti fS e lye ' s m ode l i s a dhe red t o , t h e f r equen t ly re co rded c ond i t ion o f p o ro t ic h ype ro s to s i ss hou ld n o tb er e f e r red t oa sa m a rke r o f " s t re s s " ; a l though t h e l e s ion s d o i n d ica te t h e b ody ' s a t temp t t od ea l w i th am o rb id c ond i t ion ,i nt h i sc a se t h a to fa naem ia ,t h ey d on o t r ef l ec t t h e g r ow th d i s tu rbance c onsequen t u pon h o rmona l i mba lance w h ich r e su l t s , f o r e xamp le , i nh ypop la s ia o ft h ed en ta le name l . I nc on t ra s tt oS e lye ' s i n h ib i t ing , p hy s io log ica l d e f in i t ion o fs t re s s ,t h a to fC ox a nd h i sc o l league s ( 1 983 :255 ) s e em s t o a c co rd w i th t h e g ene ra l p e r cep t ion o f s t re s s i n m o s t a n th ropo log ica l s t ud ie s :

T he d e f in i t ion s o fh ea l th w h ich w e r e o f f e r ed a t t h e b eg inn ing o f t h i s p ape r e mpha s i sed t h e d ynam ic n a tu re o f t h e c ond i t ion ; a n i n d iv idua l r a l l ies f r om i n su l t ( Audy , 1 971 ) o ra d ju s t s t oh i s o r h e r e nv i ronmen t ( Goodman , t h i s v o lume ; Wy l ie , 1 970 ) . D ubo s ( 1 965 :xv i i ) h a s d i f f e ren t ia ted b e tween a n i n d iv idua l ' s r e a c t ion t o h i so rh e re nv i ronmen t ,a nd t h er e spon se t oi t : " The c omponen t so ft h e b ody m a ch ine r e a c t w i th t h e e nv i ronmen t , i n t h e s ame w ay a sd os im i la rc omponen t si na ny o t he r l i v ing o r i n am ina te s y s tem . B u t l i v ing m an r e spond s t oh i s e nv i ronmen t . . .human a dap ta t ion c an s e ldom b e r e ga rded a s m e r e ly t h e r e su l t o f m an ' s b ody r e a c t ing w i th e nv i ronmen t a lf o r ce s " . T h i s m ay e xp la in H ink le ' s ( 1 96 1 ) o b se rva t ion o f p ro longed g ood h e a l th o n t h e p a r t o f i n d iv idua l s w ho h ave e xpe r ienced p hy s ica l , f i nanc ia l a nd e mo t iona l d ep r iva t ion , w h i le o t he r s s u ccumb t or e pea t ed i l ne s sd e sp i t el i v ing i nw ha t w ou ld s e em t o b e e x t reme ly f a vou r ab le c i r cum s tance s . D ubo s ' e mpha s i s o n t h e p e r sona l i ty o f t h e i n d iv idua l r e i te ra t e s t h e p a r t icu la r i ty o f r e spon se , a nd t h e e s sen t ia l f a c t t h a t au n ive r sa l ly " s t re s sf u l " s i tua t ion d oe s n o t e x i s t . T h i s i s e choed b y H ink le ( 1 96 1 :291 ) . S i gn i f ican t ly f o r b i o cu l tu ra l a na ly se s o f s k e le ta l r ema ins , h e s t a t e s : " Env i ronmen ta l s t r e s s ' a nd ' emo t iona l s t re s s ' s imp ly d on o te x i s ti nt h es e n se t h a tm e chan i ca l s t r e s se x i s t si n as t e e l b eam " . I n t h e f i na l a na ly s i s , t h e r e ac t ion o f i n d iv idua l s t o t h e i r s o c ia l a nd i n t e rp e r sona l e nv i ronmen t d ep end so nt h e i ro wn p e r cep t ion o ft h em . T he i n d iv idua l i ty o f r e spon se i s af a c to r o f c on s ide rab le i mpo r tance w hen m a rke r s o f i n su l t a r e i n te rp re ted i n t e rm s o f t h e h ea l th o f a c ommun i ty o r p opu la t ion a s aw ho le ( s ee B u sh a nd Z ve leb i l ,t h i sv o lume ) ,p a r t icu la r ly i nv i ew o f t h e t e ndency t o r e f e r t o " s t r e s s " i n b r oad c a tego r ie s a s d e ta i led i nT ab le 2 . 1 . I th a s b een p r opo sed , h oweve r , t h a t a l though e a ch h uman b e ing i s u n ique i n t e rms o f e xpe r ience a nd a t i tude s , t h e r e a r e m u tua l r e spon se s i n i n d iv idua l s w i th a c ommon c u l tu ra l b a ckg round ( H ink le , 1 961 ) . C on s ide r ing a ni n d iv idua lw i th in t h e f r amewo rk o f t h e i r s o c ie ty a nd t h e i r c u l tu r e m ean s t h a t t h e r e a c t ion s o ft h a t i n d iv idua l w h ich a r e p roduc t s o f c u l tu r e a nd s o c ie ty c an b e p o s tu la t ed . H ink le i n t end ed t h i s a n th ropo log ica l e v iden ce t ob e at o o l t oa i d c l in i c ian s ' e va lua t ion o f t h e i r p a t ien t s . U nfo r tuna te ly f o r a r chaeo log is t s , h i s r e commenda t ion t h a t " a

" . . the q ua l i ty o f h uman l i f e i s d ependen t o n t h e a b i l i ty t or e s i s t d i sea se , a nd t o a dap t t o t h e p hy s ica l a nd p sy cho log i ca l d emand s o f t h e e nv i ronmen t . F a i lu r et oa dap tc an r e su l ti n i mpa i r ed p hy s ica l a nd p sy cho log ica l w e l l b e ing , i nd i sea se a nd u l t ima te ly i nd ea th . T he se d emand s , t h e i r e xp e r ience , a nd e f f e c t s r e p r e sen t s t r e s s " ( empha s is i nt h e o r ig ina l ) . Y e t i tc ou ld b e a r gued , a sH ink le ( 1 96 1 ) a nd Ma son ( 1 975b ) h ave d one i nr e sp ec t o f c l in ica l w o rk , t h a t a n th ropo log ica l s t ud ie s w ou ld b ene f i t f r om t h e s t r ik ing o f t h e w o rd " s t re s s " f r om t h e v o cabu la ry , g i ven t h e p rob lems o f d e f in i t ion . E ven s o , Ma son ( 1 975b :35 ) f e l t t h a t i t s u s age i s n o t t o ta l ly d evo id o f w o r th , f o r " p e rhap s t h e n o t ion o f ag ene r ic t e rm w h ich s omehow t i e s t o ge the rt h et h r ea ten ing o rt a x ing d emand s o ft h e e nv i ronmen t o n l i v ing o r gan i sm s s t r ik e s s ome d eep , r e spon s ive c ho rd w i th in u s w h ich k eep s a l ive t h e u s e o fs t r e s s t e rm ino logy i ns p i te o fa l t h ec on fu s ion i tc r ea te s " ; p e rhap st h i se xp la in s i t s c on t inued u s e b y a n th ropo log i s t s . Mo r eove r , s eman t ic a spe c t s a s id e , i ti st h e n o t ion o f t h e o r gan i sm r e a c t ing a nd r e spond ing t oi n te rna l a nd e x ter na l a gen t s i no rde r t om a in ta in i t s i n he ren t b a lance w h ich f o rm s t h e b a s i s o f t h e c oncep t o f h ea l th u nde r ly ing t h e b i o cu l tu ra l a pp roa ch . H ea l th c anno tb ec on s ide red i n av a cuum b e cau se t h eh uman o r gan i sm i sr e cep t ive t oe nv i ronmen ta l i n f luence s , p hy s ica l , c u l tu r a l a nd s o c ie ta l ; t h e

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C ox , T . , C ox , S . & T h i r laway , M . ( 1 983 ) . T he p sycho log ica l a nd p hy s io log ica l r e spon se t o s t res s . I n P hys io log ica l C o rre la te s o f H uman B ehav iour . V o l . 1e d s . A . G a le a nd J .A . E dwa rd s . p p . 2 55 -76 . L ondon : A cadem ic P r e s s . D ubo s ,R . ( 1 959 ) . M i rage o fH ea l th . L ondon : G eo rge A l len a nd U nw in . D ubo s , R . ( 1 965 ) . Man A dap t ing . N ew H aven a nd L ondon : Y a le U n ive r s i ty P re s s . D unn , J . ( 1984 ) . S i s te r s a nd B ro the r s . L ondon : F on tana . D unn , J .a nd K end r ick , C . ( 1 982 ) . S ib l ings : L ove , E nvy a nd U nd e rs tand ing . L ondon : G ran t Mc In ty re . E ib l -E ibe s f e ld t , I .( 1 989 ) . H uman E tho logy . N ew Y o rk : A ld ine d eG ruy te r . E nge l , F .L . ( 1 956 ) . A c r i t ique o f s ome c oncep ts r e la t ing t o t h e e t iopa thogene s i s o f t h e d i seases o f a dap ta t ion . P roc . S econd Wo r ld C ong r . Med . P sycho l .B uenos A i r e s . FAO /WHO /UN ICEF P ro te in A dv i so ry G roup ( 1970 ) . L ive si nP e r i l : p ro te in a nd t h ec h i ld . R ome : FAO . G anong , W .F . ( 1 985 ) . R ev iew o f Med ica lP hy s io logy . 1 2 th e d i t ion . L o s A l to s , C a l i fo rn ia : L ange Med ica l P ub l ica t ion s . G i lb e r t , R . I .( 1 985 ) . S t re s s , p a leonu t r i t ion a nd t r ace e l emen t s . I nT he A na ly s i s o fP reh i s to r ic D ie t s . e d s . J .M ie lk e a nd R . G i lbe r t . p p . 3 39 -58 . O r lando : A cadem ic P re s s . G oodman , A .H . , A rme lago s , G .J . &R o se , J . C . ( 1980 ) . E name l h ypop la s ia s a s i n d ica to r s o f s t r e s s i n t h ree p r e -h i s to r ic p opu la t ion s f r om I l ino i s . H um . B io l . 5 2 :515 -28 . G oodman , A .H . , A rme lago s , G .J . &R o se , J . C . ( 1984 ) . T he c h rono log ica l d i s t r ibu t ion o fe name l h ypop la s ia s f r om p r eh i s to r ic D ick son Mound s p opu la t ion s . Am . J .P hys .A n th ropo l .6 5 :259 -66 . G o rdon , J . E . ( 1 969 ) . S oc ia l i mp l ica t ion s o f n u t r i t ion a nd d i sea se . A rch .E nv i ron .H ea l th / 8 :216 -34 . H ink le , L .E . ( 1 96 1) . E co log ica l o b se rva t ion s o f t h e r e la t ion o f p hy s ica l i l ne s s , m en ta l i l ne s s a nd t h e s o c ia le nv i ronmen t . P sycho soma t . Med .2 3 :289 -97 . H umn te r t , J . R . a nd V an G e rven , D .P . ( 1 983 ) . S ke le ta l g r ow th i n am ed ieva l p opu la t ion f r om S udane se N ub ia . Am .J .P hys .A n th ropo l .6 0 :471 -78 . H u s s -A shmo r e , R . , G oodman , A .H . &A rme lago s , G . J . ( 1 982 ) . N u t r i t iona l i n f e rence f r om L ev i , L . ( 1972 ) . S t re s s a nd D i sea se i n R e spon se t o P sycho soc ia l S t imu l i . O x fo rd : P e rgamon P re ss . L ev i , L . ( 1 972 ) . S t re s s a nd D i sease i nR e sponse t o P s ycho soc ia lS t imu l i . O x fo rd : P e rgamon P re ss . Mack , M .E . , B lakey , M .L . & C a r te r , S .( 1 988 ) . T he s i gn i f icance o f d en ta l d ef ec t w id th i n r e la t ion t o g r ow th a nd d eve lopmen ta l i n d ica to r s i nt h e s ke le ton ( a b s t rac t ) . Am .J .P hys .A n th ropo l . 7 5 :243 . Ma r t in , D .L . , G oodman , A .H . & A rme lago s , GJ . ( 1 985 ) . S ke le ta l p a tho log ie s a si n d ica to r s o fq ua l i ty a nd q uan t i ty o fd i e t . I n T he A na ly s i s o fP reh i s to r ic D i e t s . e d s . J .H . M ie lke a nd R . G i lb e r t . p p . 2 27 -79 . O r lando : A cadem ic P re ss . Ma son , J .W . ( 1 968 ) . A r e v iew o f p sychoendoc r ine r e sea rch o n t h e p i tu i ta ry -ad rena l c o r t ica l s y s tem . P s ycho som . Med .3 0 :576 -607 . Ma son , J .W . ( 1 97 1) . Ar e -eva lua t ion o ft h e c oncep t o f " non spec if ic i ty " i ns t re s s t h eo ry . J .P sych ia tr . R e s . 8 : 323 -33 . Ma son , J .W . ( 1 974 ) . S pec if ic i ty i nt h e o r gan iza t ion o f n eu roendoc r ine r e spon se p rof i le s . I n F ron t ier s i n N euro logy a nd N eu ro sc ience R e sea rch . e d s . P .

s o c io cu l tu ra ls y s tem c an h ave ap owe r fu li mpa c t u pon t h e i n d iv idua l , p r oduc ing p sy cho log i ca l s t r e s sw h i ch c an b em an i f e s t ed p hy s i ca l ly .

R ef erence s A rm st rong , K .F . a nd J ack son , S .M . ( 1 972 ) . A na tomy a nd P hy s io logy f o r N urse s . L ondon : B a i l l ie re T inda l l . A udy , J .R . ( 1971) . Mea su remen t a nd d i agno s i s o f h ea l th . I n E nv i ron lMe tua l e d s . P . S hepa rd a nd D . McK in ley p p . 1 40 -62 . B o s ton : H ough ton M if f l in . B oddy , J . ( 1983 ) . I n fo rma t ion p roce s s ing a nd f unc t iona l s y s tem s i n t h e b r a in . I n P hys io log ica l C o r re la te s o fH uman B ehav iou r V o l . 1 ,e d s . A . G a le a nd J .A . E dwa rd s , p p .49 -77 . O r lando : A cadem ic P re s s . B u ik s t ra , J . E . a nd C ook , D .C . ( 1 980 ) . P a laeopa tho logy : a n Ame r ican a c coun t . A nn . R ev . A n th ropo l .9 :433 -70 . C annon , W .B . ( 1914 ) . T he i n te r re la t ion s o fe mo t ion s a ss ugge s ted b yr e cen tp hy s io log ica lr e sea rche s . Am . J .P sycho ! .2 5 :256 -82 . C annon , W .B . ( 1928 ) . T he m echan i sm s o f e mo t iona l d i s tu rbance o fb od i ly f unc t ion s . N ew E ng l . J .M ed . 1 98 :877 -884 . C annon , W .B . ( 1929 ) . B od i l y C hanges i n P a in , H unge r ,F ea ra nd R age . B o s ton : C .T . B ranfo rd C o . C a r l son , D .S . , A rme lago s , G . J . & V an G e rven , D .P . ( 1 974 ) . F ac to r s i nf luenc ing t h e e t io logy o f c r ib ra o rb i ta l ia i np r eh i s to r ic N ub ia . J .H um . E vo l . 3 : 405 1 0 . C a sse l , J . , P a t r ick , R . & J enk in s , D . ( 1 960 ) . E p id em io log ica la na ly s i so ft h eh ea l th i mp l ica t ion so f c u l tu r ec hange : ac oncep tua lm od e l . A nn .N .Y .A cad . S c i .8 4 :938 -49 . C hadw ick , J .a nd Mann , W .N . ( 1950 ) . T he Med ica l Wo rk s o fH ippocra te s : an ew t r ans la t ion f rom t h e o r ig ina lG reek . O x fo rd : B la ckwe l lS c ien t if ic . C la rk , G .A . ( 1 985 ) . H e te rochrony , A l lome try a nd C ana l i za t ion i n t h e H uman V e r teb ra l C o lumn : E xamp l e sf rom P reh is to r ic Ame r ind ian P opu la t ions . P h .D . D i s se r ta t ion , U n ive r s i ty o f Ma ssachu se t t s , Amhe r s t . A nn A rbo r : U n ive r s i ty M ic rof i lm s . C la rke , S .K . ( 1978 ) . Ma rke r so f Me tabo l ic I n su l t : t h e A s so c ia t ion o fR ad iopaque T ransve r se L ine s , E name l H ypop la s ias a nd E name l H i s topa tho log ies i n a P reh i s to r ic H uman S ke le ta l S amp le . P h .D . T he s i s . U n ive r s i ty o f C o lo rado , B ou lde r . A nn A rbo r : U n ive r s i ty M ic rof i lm s . C ohen , M .N . a nd A rme lago s , G .J . ( 1 984 ) . e d s . P a leopa tho logy a t t h e O r ig ins o f A g r icu l tu re . O r lando : A cadem ic . C ondon , K .W . ( 1981) . C o r respond ence o f D eve lopmen ta l E name l D ef ec t s b e tween t h e Mand ibu lar C an ine a nd F i rs t P remo la r . U npub l i shed M .A . D i s se r ta t ion , D epa r tmen t o f A n th ropo logy ,U n ive r s i ty o fA rkan sa s ,F aye t t ev i l le . C ook , D .C . ( 1979 ) . S ub s i s tenc e b a se a nd h ea l th i n p r eh i s to r ic I l ino i s V a l ley : e v idence f r om t h e h uman s ke le ton . Med .A n thropo l .4 : 109 -24 . C ook , D .C . ( 1 981) . Mo r ta l i ty , a ge s t ruc tu re a nd s t a tu s i nt h e i n te rp re ta t ion o fs t re s s i n d ica to r s i np r eh i s to r ic s k e le ton s : ad en ta l e xamp le f r om t h e L owe r I l ino i s V a l ley . I n T he A rchaeo logy o f D ea th . e d s . R . C hapman , I .K inne s & K . R and sbo rg . p p .133 -44 . C amb r idge : C amb r idge Un ive r s i ty P re s s . C ox , T . ( 1978 ) . S t res s . L ondon : MacM i l lan P re s s L td .

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R o se , J . C . a nd P a s ley , J .N . ( 1980 ) . S t res s a nd d en ta l d eve lopmen t : a n e xpe r imen ta l p a leopa tho log ica l m ode l . P ape r p r e sen ted a t4 9 th A nnua l Mee t ing o f t h e Ame r ican A s soc ia t ion o f P hy s ica l A n th ro p o log i s t s , N iaga ra F a l l s ,N ew Y o rk . R o se , LC . , C ondon , K .W . & G oodman , A .H . ( 1 985 ) . D ie t a nd d en t i t ion : d eve lopmen ta l d i s tu rbance s . I n T he A na l ys i s o fP reh is to r ic D ie ts e d s . J .H . M ie lke a nd R . G i lb e r t . p p . 2 81-305 . O r lando : A cadem ic P r e s s . R udney , J .( 1981) . T he P a l eoep idem io logy o fE a r l y C h i ldhood S t re s s i n T wo A nc ien t N ub ian P opu la t ions . P h .D . d i sse r ta t ion , U n ive r s i ty o f C o lo rado , B ou lde r . S c r im shaw , N .S . , H ab ich t , J . P . , P i che , M .L . , C ho lako s , B . & A r royave , G . ( 1 966 ) . P ro te in m e tabo l i sm o f y oung m en d u r ing u n ive r s i ty e xam ina t ion s . Am . J . C l in .N u t r ./ 8 :321 -24 . S e lye , H . ( 1 950 ) S t res s . Mon t rea l : A c ta I n c . Med . P ub l . S e lye , H . ( 1 957 ) . T he S t re ss o f L if e . L ondon : L ongman s ,G reen a nd C o . S e lye , H .( 1 973 ) . T he e vo lu t ion o ft h e s t r e s s c oncep t . Am .S c i . 6 1 :692 -99 . S e lye , H . ( 1 976 ) . F o r ty y ea r s o f s t re s s r e sea r ch : p r inc ipa l r ema in ing p rob lem s a nd m i sconcep t ion s . C an . Med .A s soc .J . 15 :53 -6 . W iddow son , E .M . ( 1951) . Men ta l c on ten tmen t a nd p hy s ica lg r ow th . L ance t1 : 1316 -18 . Wy l ie , C .M . ( 1 970 ) . T he d ef in i t ion a nd m ea su remen t o fh ea l th a nd d i sea se . P ub .H l th .R ep .8 5 :10 0 -04 .

S eeman a nd 0 . B rown . p p . 6 8 -80 . T o ron to : U n ive r s i ty o fT o ron to . Ma son , J .W . ( 1 975a ) . Ah i s to r ica l v i ew o f t h e s t re s s f i e ld . P a r tI .J .H um .S t re s s . / : 6 -12 . Ma son , J .W . ( 1 975 1 , ) . Ah i s to r ica l v i ew o f t h e s t re s s f i e ld . P a r tI . J .H uman S t re s s . 2 2 -36 . Men sfor th ,R .P . ( 1 986 ) . T he p a thogene s i so fp e r io s t ea l r e a c t ions i n e a r l ie r h uman g roup s : d i agno s i s , e p idem io log ica l a nd d emog raph ic c on s ide ra t ion s ( a b s t rac t ) . Am .J .P hys .A n th ropo l . 6 9 :239 . Men sfo r th , R .P . , L ove joy , C .O . , L a l lo , J .W . & A rme lago s , G . J . ( 1978 ) . T he r o le o f c on s t i tu t iona l f a c to r s , d i e t a nd i n f ec t iou s d i sea se i nt h e e t io logy o f p o ro t ic h yp e ro s to s i s a nd p e r io s tea l r e a c t ion s i n p r eh i s to r ic i n fan t s a nd c h i ld ren . Med . A n thropo l . 2 : 1 -59 . Mo s s , G .E . ( 1 973 ) . I lne s s , I mmun i ty a nd S oc ia l I n te rac t ion . N ew Y o rk : J ohn W i ley a nd S on s . P a lkov ich , A .M . ( 1 987 ) . E nd em ic d i sea se p a t t e rn s i n p a laeopa tho logy : p o ro t ic h ype ro s to s i s . Am .J .P hys . A n thropo l . 7 4 :527 -37 . P a rk , E .A . ( 1954 ) . B one g r ow th i nh ea l th a nd d i sea se . A rch .D is .C h i ld .2 9 :268 -81 . P o r te r , R . ( 1987 ) . D i sea se , Med ic ine a nd S oc ie ty i n E ng land 1 550 -1860 . B a s ing s toke : Ma cm i l lan E duca t ion . P owe l l , G .F . , B ra se l , J .A . & B l izza rd , R .M . ( 1 967 ) . Emo t iona l d ep r iva t ion a nd g row th r e ta rda t ion s imu la t ing i d iopa th ic h ypop i tu i ta r i sm . I . C l in ica l e va lua t ion o f t h e s ynd rome . N ew . E ng . J . Med . 2 76 :1271-78 . P owe l l , G .F . , B ra se l , J .A . & B l izza rd , R .M . ( 1 967 ) . Emo t iona l d ep r iva t ion a nd g row th r e ta rda t ion s imu la t ing i d iopa th ic h ypop i tu i ta r i sm . E ndoc r ino log ic e va lua t ion o f t h e s ynd rome . N ew . E ng .J . Med .2 76 :1279 -83 . P owe l l , M .L . ( 1 985 ) . H ea l th , D i sea se a nd S oc ia l O rgan i za t ion i n t h e M i s s i s s ipp ian C omn zun i ty a t Moundv i l le . P h .D . d i s se r ta t ion . N o r thwe s te rn U n ive r s i ty , I l ino i s . R o se , J .C . ( 1 977 ) . D ef ec t ive e name l h i s to logy o f p r eh i s to r ic t e e th f r om I l ino i s . Am . J . P hys . A n thropo l .4 6 :439 -66 . R o se , J .C . a nd B oyd , L . ( 1 978 ) . D ie ta ry r e con s t ruc t ion u t i l iz ing h i s to log ica l o b se rva t ion o f t h e e name l a nd d en t in ( ab s t rac t ) . A n z .J .P hy s .A n thropo l .4 8 :431 .

A cknow ledgemen ts T he d o c to ra lr e sea r ch f r om w h ich t h i sp ape ri s r e v i sed w a s f u nded b y t h e S c ience a nd E ng inee r ing R e sea r ch C ounc i l . It h ank D r . T . C ox , D epa r tmen t o f P sy cho logy , U n ive r s i ty o f N o t t ingham , P ro f . R .G .G . R u s se l l , D epa r tmen to f H uman Me tabo l i sm a nd C l in i ca l B iochem i s t ry , R oya l H a l lamsh i r e H o sp i ta l , S he f f ie ld a nd P ro f . J .M . T anne r , I n s t i tu te o fC h i ld H ea l th , U n ive r s i ty o f L ondon , f o r t h e i r h e lp f u l c ommen ts o n a n e a r l ie rv e r s ion . Ia m g r a te fu l t oD r . P e te r Sm i th , R eade ri nP sycho logy ,U n ive r s i ty o fS he f f ie ld ,f o r u s e f u ld i s cu s s ion .

1 9

T ab l e2 . 1 .

T he a n th ropo log i ca lu s age o f" s t r e s s " .

U sage

P ub l ica t ion

A s an on spe c if ic p hy s io log ica ls t a te :

C ondon , G oodman e ta l , G oodman e ta l , H u s s -A shmo r ee ta l , R o se a nd B oyd , R o se a nd P a s ley , R o s ee ta l ,

" ma lnu t r i t ion ,d i sea se a nd o t he r p hy s io log ica ls t r e s se s "

1 98 1 1 980 1 984 1 982 1 9 79 1 980 1 985

H umme r ta nd V an G e rv en , 1983 :47 1 H umme r ta nd V an G e rven , 1 9 82 :1

" b io log ica ls t re s s " " s y s tem ic s t r e s sf r om s p e c i f ic o r n on spec i f ic i n f e c t ion a nd /o rf r om g ene r a lo rs p ec if ic n u t r i t iona l d e f ic ienc ie s "

P owe l l , 1 985 :203

" . . r e spon se t on u t r i t iona ls t r e s s "

M en s fo r th e ta l , 1 982 :1

" c h ron ic o re ndem i cd i e ta ry s t r e s s "

P a lkov i ch , 1 987 :527 .

" i n f ec t iou sd i sea se s t r e s s "

Men s fo r th , 1 986 :239

" c h i ldhood s t re s sa nd m o rb id i ty "

H umme r ta nd V an G e rven , 1 983 :47 1

" t he e f f e c t so fs t r e s st h a td en ta l e name ld e f e c t sr e p r e sen t "

Ma ck e ta l , 1 988 :243

" ah uman p opu la t ion u nde rs t r e s s "

M en s fo r th e ta l , 1 982 :2

" h igh s t a tu si n d iv idua ls e xp e r ienced l e s ss t re s s "

C ook , 1 98 1 :140

" p sycho soc ia ls t r e s s "

G i lb e r t , 1 985 :341

2 0

2 1

T he G ene ra lA dap ta t ion S ynd rome ( f rom S e lye , 1 957 ) .

R es is tance t o s tress

I .

b i ) 4 .

3 .

ACCOUNTING FOR THE EVOLUT ION OF P SYCHO SO MAT IC PHENO MENA : D ID UNCO -OPERAT IVE UPPER PALAEOL ITH IC P EOPLE BECO ME I LL AND D IE?

M ichae lE .H y land a nd W in S cu t t . f o l low c hange s i n m o rpho logy ( e .g . i n c rea se o f s k u l l c apa c i ty ) , o rs ome f o rm o fc u l tu ra l c hange w h ich e nhance s t h ink ing s k i l l s ( e .g . e a r ly i n fan t s t imu la t ion ) . T h ink ing s k i l l s a r e t h ough t n o t t o i mp rove e xcep t w i th m o rpho log ica l a nd c u l tu ra l c hange . A r e la ted a s sump t ion i s t h a t s im i la r m o rpho log ie sa r ei n d i ca t ive o fs im i la rg eno type s . T he f a i lu r e t o r e cogn i se t h a t p sycho log ica l c apac i t ie s m ay h ave a g ene t i c c omponen t i sa d i f f e r en ce b e tween a r chaeo logy a nd p sycho logy . C e r ta in ly i ti st h eo r e t ica l ly p o s s ib le t h a t s im i la r m o rpho log ie s m ay b e a c compan ied b y d i f f e r en t g ene t i ca l ly d e te rm ined p sy cho log ie s . H oweve r , a n e xamp le o f t h e w eakne s s o f r e ly ing o n m o rpho log ica l s im i la r i ty t o i n f e r t o ta l g ene t ic s im i la r i ty i sp r ov ided b y t h eb r own t r ou t a nd s e a t r ou t ( t he l a t te r i sd e sc r ibed b y a ng le r s a s a" s ea r u n b r own t r ou t " ) w h i ch h ave t h e s ame m o rpho logy a nd s o a r e b o th c l a s s i f ied a s S a lmo t r u t ta b u t h ave d i f f e r en t b ehav iou r p a t ter n s . E v idence f o r g ene t ic d i s s im i la r i ty b e tween s e a a nd b r own t r ou t i s p r ov id ed b y s mo l t t r an s fo rma t ion w h ich o ccu r s i n s e a t r ou t e ven w hen r a i sed i nf r e shwa te r h a tche r ie s b u t n eve ri n b r own t r ou t . I ti sn ow t h ough t t h a t t h e b rown t r ou t e vo lved f r om t h e s e a t r ou t a s p a r t o f t h e r e co lon iza t ion o f r i ve r s a f te r t h e l a s t i c e a ge ( Ha r r i sa nd Mo rgan , 1 989 ) . T he s u cce s s o f as p ec ie s d epend s n o t o n ly o n i t s m o rpho logy b u t a l so o n i t sb ehav iou r , a nd s o p sy cho log i ca l e vo lu t ion m u s t b e r e levan t t o a n i n te rp r e ta t ion o ft h e a r cha eo log i ca l r e co rd . T hu s f o ri n s tance ,t h ee a r ly H omo s ap ien ss ap iens f r om Q a f zeh a nd S khu l w ho h ave n ow b een d a t ed t o a bou t 9 0 ,000 B P a nd w ho se l i t h ic a r t i f ac t s r e p r e sen t a L eva l lo i s -Mou s te r ian ( M idd le P a la eo l i th ic ) i n du s t ry ( S t r inge r , 1 988 ; S t r inge r , G r un , S chwa r cz & G o ldbe rg , 1 989 ) m ay n o t n e ce s sa r i ly r e p re sen t t h e s ame g ene p oo l a s a na tom ica l ly m ode rn h uman s d a t ing f r om t h e U pp e r P a laeo l i th ic . A na tom ica l s im i la r i ty i sa n e ce s sa ry b u t n o t a s u f f i c ien t c ond i t ion f o r g ene t i cs im i la r i ty ,a nd ac hange i nb ehav iou rw i th m o rpho log ica l c on s tancy m ay n o t b e e xc lu s ive ly t h e r e su l t o f e x t r a soma t ic a dap ta t ion b u t m ay r e f lec t s ome i n t ra soma t ic a dap ta t ion i nt h e f o rm o f ac hange t ot h eg ene t ic s ub s t ra t eo fb ehav iou ra l c apa c i t ie s . T he p u rpo se o f t h i s p ape r i st oe xp lo r e o ne p a r t icu la r i n te r f a ce b e tween p sy cho logy a nd a r chaeo logy . I nr e cen t y ea r s , t h e re h a s b een a n a c cumu la t ion o f e v idence t h a t p sycho log ica l s t a te s h ave a c a u sa l r e la t ion sh ip w i th m o rb id i ty

T he r e la t ion sh ip b e tween a r chaeo logy a nd p s ycho logy h a s r e ce ived l i t t le a t t en t ion f r om e i the r a r cha eo log i s t s o r p sycho log i s t s . T he t h eo re t ica lp o t en t ia lo fr e sea r ch a tt h ei n t e r f a ce o f t h e se t wo d i sc ip l ine si so f t en o b s cu r ed ,a tl e a s ti n p a r t , d ue t o as imp l i f ied v i ew m embe r s o fe i the r d i sc ip l ine h ave o ft h eo t he rd i sc ip l ine . T o a p sycho log i s t , b ehav iou r i s t h e c on sequence o f a n i n te ra c t ion b e tween g ene t ic a nd e nv i ronmen ta l f a c to r s . T he w ay g ene t ic a nd e nv i ronmen ta l f a c to r s i n te ra c t t o p r oduce b ehav iou rv a r ie sa l ong t h ep hy logene t i c s c a le . I n t h e c a se o f s imp le c r ea tu r e s , s u ch a s s p id e r s o r f r ogs , b ehav iou ra l p a t t er n s a r e g ene t i ca l ly d e te rm ined ( a nd r e f e r r ed t o a s " f ixed a c t ion p a t te rns " ) . I nt h ec a se o fh uman s ,w he r et h e r ea r e f ew f i xed a c t ion p a t ter n s , t h eg ene t ic c on t r ibu t ion i sp r ima r i ly o ne o f ac apa c i ty , a nd b ehav iou r i s t h ec onsequence o ft h ei n t e r a c t ion o ft h a tc apa c i ty w i th e nv i ronmen ta l c ond i t ion s ( Ba te son , 1 979 ) . F o r e xamp le , h uman s , b u t n o t d og s , h ave t h e g ene t ica l ly t r an sm i t ted c apa c i ty t ol e a rn l a nguage g i ven t h er i gh te nv i ronmen t a lc ond i t ion s . F o r m any p sycho log i s t s , t h e g ene t i c c on t r ibu t ion t o b ehav iou r i s a s sumed t o h ave e vo lved b ecau se i tw a sb i o log i ca l ly a dap t ive ,t h a t i s ,a dvan tageou st ot h eg ene p oo l . B u tc on t ra ry t o e c o log ica l t h eo ry , t h e c oncep t o f b i o log i ca l a dap ta t ion i s d i vo r ced f r om t h e e co log ica l e nv i ronmen t . F o r e xamp le , L o r enz ( 1 966 ; s e e a l so B a i ley , 1 987 ) s ugge s t s t h a ta gg re s s ion h a s a g ene t ic b a s is b e cau se i ti sb i o log i ca l ly a dap t ive ; o n t h e o t he r h and , R u sh ton , R u s se l l a nd We l l s ( 1 984 ) s ugge s t t h a t a l t ru i sm h a s ag ene t i c b a s i s b e cause i t i s b i o log ica l ly a dap t ive . B o th p s y cho log i s t s f a i l t o r e cogn i se t h a t w ha t i s b i o log ica l ly a dap t ive i n o ne e co log ica l e nv i ronmen t m ay n o t b e b i o log i ca l ly a dap t ive i n a no the r . I n s ho r t , t h e e vo lu t ion o f t h e g ene t ic c on t r ibu t ion t o b ehav iou r c a nno t b e u nde r s tood d i vo r ced f r om i t s e nv i ronmen ta l c on t ex t . I tc a n b e u nd e r s tood o n ly i n t e rms o f as e r ie s o f e vo lu t iona ry e nv i ronmen t s a s p r e sen ted b y t h e a r chaeo log ica l r e co rd . T o b e c on s i s ten t w i th e vo lu t iona ry t h eo ry , t h e p sy cho log i s t s ' s c oncep t o ft h e b i o log ica l u t i l i ty o fb ehav iou r w i l l h ave t o d epend o na r chaeo logy . T o mo s t a r chaeo log i s t s , a n o r gan i sm ' s b ehav iou r i s a s sumed t o b e d i c ta ted b y i t s p hy s ica l f o rm a nd i t s c u l tu r a l o r " e x t ra soma t i c " ( Wh i te , 1 959 ) m ean s o fa dap ta t ion . T hu s , i nt h e c a se o f h uman s , p sycho log ica l c hange s ( e .g . , i mp rovemen t i n t h ink ing s k i l l s ) a r e a s sumed t o

2 3

( i l lne s s ) a nd d ea th ( mo r ta l i ty ) . F o l low ing e a r ly e v idence t h a t b e reavemen t i n c r ea se s mo r ta l i ty i n t h e r ema in ing s pou se , t h e r e n ow s e em s t o b e c on s id e rab le ( t hough b y n o m ean s u ncon t rove r s ia l ) e v idence t h a t p sycho log ica l f a c to r s p l ay ar o le i nt h e i n c idence a nd r e cove ry o fa l l p hy s i ca l d i sea se -s u ch a s r e sp i ra to ry t r a c t i n f e c t ion s , c o ld so r e s , c ance r , h ea r t d i sea se , a r th r i t i s a nd s o o n . C on t r a ry t ot h e a s sump t ion s o f t r ad i t iona l p hy s ica l m ed i c ine ( r e f e r r ed t o i n m ed ic ine a s " t he m ed ica l mode l " ) - a mode l r e f lec ted i n s ome o f t h e o t he r p ape r s i n t h i s v o lume -t h e i n c iden ce o fp hy s ica l i l ne s s i sn o t t h e r e su l t o f o n ly d i e t a nd o t he r p hy s i ca l c ond i t ion s . T he i n c id ence o f p hy s i ca l i l ne s s i s a f f e c t ed b y m any c au se s , o ne o f w h ich i s t h e p sy cho log i ca l s t a te o f t h e i n d iv idua l . I ti sn ow r e cogn i sed ( Bake r , 1 987 ) t h a t t h e t e rm p sy cho soma t ic i l ne s s s hou ld a pp ly n o tt o as ma l l n umbe ro fi l ne s se sb u tc an b ec ha ra c t e r i s t ic o fa l l p hy s ica li l ne s s . I f t h e r e i s a r e la t ion sh ip b e twe en p sy cho log i ca l s t a te s a nd p hy s ica l i l ne s s , t h i s r e la t ion sh ip h a s e i the r a r i sen b y c hance o r i th a s e vo lved . T he p u rpo se o f t h i s p ape r i s t o d emon s t ra te t h a t t h e r e la t ion sh ip b e tween p sycho log ica l s t a te s a nd m o rb id i ty , h ence fo r th r e f e r red t oa sp sycho soma t ic p henom ena , d i d n o t e vo lve b y c hance ; i te vo lv ed t h rough n o rma l e vo lu t iona ry p roce s se s a nd t h e p e r iod o f e vo lu t ion c an b e u nde r s tood f r om ac omb ina t ion o fe x i s t ing a r chaeo log ica la nd p sycho log ica ld a ta .

p a r en t ' s r o t t ing c a r ca se p rov ide s ar i ch s o u r ce o f f o od . Wha t i ss i gn if i can t a bou t t h e d ea th o f t h e s o ckeye s a lmon i st h a t s ene scence i sc on t ro l led n o t b y a n " i n t e rna l c l o ck " o rs ome o t he r i n te rna l , p hy s io log ica l m e chan i sm b u t t h rough a b ehav iou ra l t r i gge r . T he b ehav iou r o f s p awn ing t r igge r s o f f a n i n te rna l s e l f -de s t ruc t p ro ce s s w h i ch t a ke st h ef o rm o fa c ce le ra ted s ene s cence . I th a s l o ng b e en r e cogn i sed t h a t p r e sene s cen t d ea th c an b e b i o log i ca l ly a dap t ive . T he c l a s s ic e xamp le o ft h i sp henomenon i st h ec a se o fw o rke r b ee s s a c r if ic ing t h em se lve s t os ave a h i ve u nde r a t ta ck . T he c au se o f d ea th w hen w o rk e r b ee s s a cr i f ice t h em se lve s i se x t erna l t ot h e b ee ( i . e . , i t i st h e a t ta cke r ) . H oweve r , i ti sa l so p o s s ib le f o r p r e sene s cen t d ea th t o b e b io log ica l ly a dap t ive w he r e t h e r e i sa n i n te rna l m e chan i sm f o r d ea th , a s i n t h e c a se o f p o s t rep roduc t ive s o ckeye s a lmon . T he r e a r e t wo w ay s i nw h ich i n t e rna l ly d r iven , p r e sene scen t d ea th c an b e b i o log i ca l ly a dap t ive , t h e s e t wo w ay s o ccu r r ing i ne i th e r aK o rr env i ronmen t( P ianka , 1 988 ) . A K env i ronmen t i s c ha r a c te r ised b y h i gh i n t ra sp ec if i c c ompe t i t ion f o r s ca rce r e sou r ce s , s u ch t h a t p opu la t ion g r ow th i s c on s t ra ined b y p opu la t ion s i z e . U nde rs u ch c ond i t ion s ,i tm ay b e b i o log ica l ly a dap t ive i fw eake r m embe r s o f t h e s p e c ie s b ecome i la nd d i e , a s b y d y ing t h ey d o n o t c ompe t e w i th t h e s upe r io r m embe r s o f t h e s p e c ie s f o r s c a r ce r e sou r ce s . E ndo c r ino log ica l c hange s i ns o c ia l ly i n f e r io r m embe r s o f as p ec ie s i s a r e cogn i sed f o rm o f p opu la t ion c on t ro l ( Ch r i s t ian , 1 980 ) . A g ood e xamp le o f t h is p henomenon i sp rov ided b y t r ou t w ho a r e s o c ia l a n ima l s a nd h ave d om inance h i e ra r ch ie s . T rou t a t t h e b o t tom o f t h e h i e ra r chy t e nd t o b e i mmuno supp r e s sed ( i . e . , h ave l e s s a c t ive i mmune s y s tems ) a nd a r e t h e re fo r e mo r e l i ke ly t o s u ccumb t oi n f ec t ion s( P e te r se ta l , 1 988 ) . A n r env i ronmen t i s c ha ra c t e r i s ed b y r e la t ive ly l ow i n t ra spe c i f ic c ompe t i t ion f o rs c a r ce r e sou r ce s s u ch t h a t p opu la t ion g r ow th i s u ncon s t ra ined b yp opu la t ion s i ze . I ft h e r e w e r e a f o rm o fr env i ronmen t w he r e t h e s u rv iva l o f t h e s pe c ie s d ep end s c r uc ia l ly o n c o -ope ra t ion b e twe en g r oup m emb e r s , t h en , u nde r s u ch c ond i t ion s , u nco -ope ra t ive m embe r s c an t h r ea t en t h e s u rv iva l o f t h e i r g r oup a nd t h e r eby t h r ea ten s u rv iva l o f s im i la r g ene t ic m a te r ia l . I tw ou ld t h e r e fo r e b e b i o log ica l ly a dap t ive i fs u ch u n co o p e ra t ive i n d iv idua l sb ecame i la nd d i ed a st od o s o w ou ld e nhance t h e s u r v iva l c hance s o fo t he r s a nd o ft h eg ene p oo l .

Two Way si n Wh ich P re sene scen t Mor ta l i ty i sB io log ica l ly A dap t ive I n o u r s a nd m any o t he r c u l tu r e s , d ea th i s v i ewed s omewha t n ega t ive ly . T h i s n ega t ive a pp roach t o d ea th c ome s a bou t f r om o u r o wn i n d iv idua l i s t ic p e r spec t ive : p eop le t r yt op r e se rve t h e i ro wn l i ve s a nd t h o se o ft h e i r l o ved o ne s . A t t h e l e ve l o f t h e i n d iv idua l , t h e d ea th o f ac l o se f r i end o r r e la t ive i sa lway s at r agedy . T he r e i s , h oweve r , a n a l ter na t ive a nd m uch mo re p o s i t ive v i ew o f d ea th w h i ch c ome s f r om a g ene t i c p e r spec t ive . I n t h e c a se o f s exua l r e p roduc t ion , m a in tenance o f t h e g ene t ic m a te r ia l ( t he p u rpo se o f t h e g ene ) i sa ch ieved t h rough t h e c r ea t ion o f n ew o r gan i sm s w h i ch a r e s l igh t ly d i f f e r en t f r om t h e i rp a r en t s ,a nd f rom t h ed ea th o ft h o se p a ren t s . F rom a g ene t ic p e r sp e c t ive , d ea th , a nd i n p a r t icu la r , d ea th f r om s e ne s cence , i sa n e s sen t ia l p a r t o f b i o log ica l a dap ta t ion , w he r e b i o log ica l a dap ta t ion i sd ef ined a s ac ha r a c te r i s t i c w h ich e nhance st h ec on t inua t ion o ft h eg ene p oo l . N o t o n ly i s s e ne s cen t d ea th b i o log ica l ly a dap t ive ,b u ti ns ome c a se st h ed ea th o fp a ren t si s e xqu i s i te ly t i med t o i mp rove b i o log ica l a dap ta t ion o f t h e s pe c ie s . T he s o ckeye s a lmon s pawn sa nd d i e s w i th in ac oup le o fd ay st h ough a p r oce s s o f a c ce le ra ted s e ne s cence ( Cou s t eau , 1 975 ) . B y d y ing , t h e p a r en t s o ckeye s a lmon d o n o tc ompe t e f o rf ood w i th t h e i ro f f sp r ing ,a nd t h e

E vo lu t ionary F orce si nF ood C o l lec t ing a nd F ood P roduc ing S oc ie t ie s F o rH omo s ap ien s ,t h eU ppe rP a la eo l i th ic w a s a n r env i ronmen t c ha r a c t e r i sed b y d i sp e r sa l i n to n ew r e g ion s ( Gamb le , 1 986 ; G e i s t , 1 978 ) . G lac ia t ion , h i gh w ind s , a nd r educed r a in f a l l d u r ing t h a t p e r iod w ou ld h ave i n t rodu ced e nv i ronmen ta l s t r e s s w h ich c ou ld h ave b een

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H oweve r , a l though N eo l i th ic s o c ie ty d epended p a r t ly o n c o -ope ra t ion , c o o p e ra t ivene s s n o l o nge r h ad t h e b i o log ica l u t i l i ty i th ad h ad i nt h e P a laeo l i th ic , a nd t h e r e a son f o r t h i sc hange r e la te st og roup s i ze . I f ap e r son w a s u nco -op e r a t ive i n a P a laeo l i th ic g roup , t h e r e la t ive ly s ma l l s i ze o ft h eg roup w ou ld m ean t h a t t h e u nco -op e ra t ive p e r son c ou ld a f f ec t t h e w o rk ing o f t h e g roup a s aw ho le , a nd h ence t h e g r oup ' s s u rv iva l . S u rv iva l w a s s o p r eca r iou s i n t h e U ppe r P a la eo l i th ic t h a t u n co -ope ra t ivene s s w ou ld h ave a f f ec ted t h e g ene p oo l . N o s o i nt h e N eo l i th ic , w he r e t h e i n c rea sed p opu la t ion s i ze w ou ld h ave a c ted a s a b uf f e r a ga in s t a n i n d iv idua l ' s u n so c ia l a c t s . T hu s a l though t h e d eve lopmen t o f aN eo l i th ic c ommun i ty r equ i r e s c o -ope ra t ion , o nce t h a t c ommun i ty h a s d eve loped , c o -op e ra t ivene s s h a s l e s s b i o log ica l u t i l i ty . Mo reove r , t h e i n c r ea sed p opu la t ion d ens i ty i n f o od p roduc ing a nd p a r t icu la r ly u rban s o c i e t ie s w ou ld h ave s h i f ted t h e e nv i ronmen t i n to a K e nv i ronmen t . A s p opu la t ion d en s i ty i n c rea sed , w i th in spe c ie s c ompe t i t ion f o r s c a r ce r e sou r ce s w ou ld h ave e nhanced t h e b i o log ica l u t i l i ty o f c ompe t i t ivene s s . E v idence s how s t h e c hange f r om a nr env i ronmen tt o aK -env i ronmen tt ob ea g r adua l o ne ( Bende r , 1 975 ) . T o t a ke t h e l a te r P reh i s to ry o f E u rope a s a n e xamp le , c ompe t i t ivene s s c an b e s e en i n t h e s h if t f r om g ene ra l i sed t on ega t ive r e c ip ro c i ty ( Sah l in s , 1 968 ; S e rv i ce , 1 966 ) , t h e g row th o f e xc lu s ive t e r r i to r ia l i ty , n ume ra cy a nd w ea l th ( Renf r ew , 1 972 ) a nd i nt h e i n c r ea s ing i n c idence o fa r t if ac t s r ef l ec t ing w a r f a r e , f r om w eapon s i n h om ic ide c on t ex t s t o f o r t i f ica t ions ( Newe l l , 1 984 ; Me ik le john e ta l , 1 984 ) . T he b i o log ica l u t i l i ty o f c omp e t i t ion i s e v iden t i n r e cen t t i me s w he r e s e t t le r s i nN o r th Ame r ica a nd A us t ra l ia c ompe ted s o s u cce s s f u l ly w i th t h e i n d igenou s p opu la t ion s t h a t t h ey h ave p ro spe r ed a t t h e e xpen se o f t h e i n d igenou s p opu la t ion s . T hu s , f r om a n e vo lu t iona ry p e r spec t ive , f ood p roduc ing a nd u r ban s o c ie t ie sa r ec ha ra c te r i sed b y as t rong f o r ce t owa rd s i n t ra sp ec i f ic c omp e t i t ion i n t h e s p ec ie s H omo s ap iens w h i ch f r equen t ly t a ke s t h e f o rm o f r e g iona lc onf l ic t .

a lev ia ted t o s ome d eg r ee t h rough c o -ope r a t ion . T he re i s c on s ide rab le e v idence t h a t U pp e r P a laeo l i th ic p eop le ( Homo s ap iens s ap iens ) a nd t o al e sse r e x ten t N eand e r tha l s ( Homo s ap ien s n eand er tha lens is ) w e r e h i gh ly c o -ope ra t ive i n t h e i r g a the r e r /hun te r / f i sh e r o r f ood c o l lec t ing e conomy ( Gamb le , 1 986 ; G e i s t , 1 978 ) . B o th l i ved i n s o c ia l g roup s , b o th e ngaged i n c o o p e ra t ive h un t ing , b o th c a r ed f o r t h e s i ck a nd b o th u sed s ymbo l i c b u r ia l o f t h e d ead . T he r e i s c i r cums tan t ia l e v iden ce t h a t c o -ope ra t ive s k i l l s w e r e p a r t icu la r ly d eve lop ed i nt h e c a se o fU ppe r P a laeo l i th ic p eop le : c amp s i te s w e r e l a rge r t h an i nt h ec a se o ft h eN eande r tha l s ; h om i c ide w a sl e s s f r equen t i n N eo l i th ic g r oup s ; a nd t h e w ide d i s t r ibu t ion o f a r t i f a c t s s ugge s t s t h a t c r o s s r e g iona l c o -ope ra t ion t o ok p l ace ( Ge i s t , 1 978 ) . E x t rapo la t ion s f r om m oder n d ay g a the r e r /hun te r / f ishe r s s ugge s t t h a tH omo s ap iens s u rv ived i n aP a laeo l i th ic e nv i ronmen t t h rough a c omb ina t ion o f i n t r ag roup ( l oca l ) a nd i n te rg roup ( r eg iona l ) c o -op e r a t ion . F o r i n s tance , B in fo rd ( 1 983 ) s how s t h a t i f t h e re i s ac a ta s t roph ic d ec rea se i nf o od r e sou r ce s i na n a r ea o ccup ied b y o ne I nu i t g r oup , t h a t g r oup w i l l m ove a s g ue s t s i n to a no the rg roup ' s a r ea . S uch c o -ope ra t ion i sa u s e fu l s t r a tegy f o r s u rv iva l o f as p ec ie s w hen s u rv iva l i ss o p r eca r iou s . F rom a n e vo lu t iona ry p e r spe c t ive , t h e P a laeo l i th ic , a nd p a r t i cu la r ly p e r iod s o f g l ac ia t ion d u r ing t h e P a laeo l i th i c , w a s a n e nv i ronmen t w i th as t rong e vo lu t iona ry f o r ce f o rc o -ope ra t ion w i th in t h e s p ec ie sH omo s ap ien s ( B in fo rd , 1 983 ;G amb le , 1 9 86 ) . T he mo r e b en ign c l ima te w h i ch m a rk s t h e o n se t o ft h e Me so l i th i c i n i t ia ted ag r adua l s h i f tt o aN eo l i th ic , p r ima r i ly f ood p rodu c ing e conomy b a sed o n a g r icu l tu r e a nd a n ima l h u sband ry . T he c hange f r om f ood c o l le c t ing t o f o od p r oduc ing h ad a p ro found e f f ec t o n p opu la t ion d en s i ty . N eo l i th ic c ommun i t ie s c ha r a c te r i s t i ca l ly h ave a n i n c r ea sed p opu la t ion d en s i ty a nd a gg r ega t ion s o f g r ea te r s i ze t h an e i the rP a la eo l i th ic o r Me so l i th i c c ommun i t ie s . T he r e a son w hy N eo l i th ic c ommun i t ie s d eve loped r e la t ive ly s imu l taneou s ly a r ound t h e w o r ld d u r ing t h e p r e sen t i n t e rg la c ia l r a the r t h an d u r ing a n e a r l ie r i n te rg la c ia l h a s b een t h e s ub jec t o fm uch c on t rove r sy . T ot h o se o t he re xp lana t ion s ( s uch a s p ack ing , e . g . B in fo rd , 1 983 ; s e e a l so B end er , 1 975 ) w e n ow a dd a p sycho log ica l e xp lana t ion . T he d ev e lopmen t o f aN eo l i th i c c ommun i ty w ou ld h ave r e qu i r ed t h e p sy cho log i ca lc apa c i ty f o ra dvanced c o -op e ra t ion . T ha t i s , i tw ou ld b e n e ce s sa ry f o rp eop le t oh ave b een s k i l led i nc o -oper a t ion . I ft h e p sycho log ica l c apac i ty f o r c o -ope ra t ion w a s a n U pp e r P a la eo l i th ic e vo lu t iona ry d eve lopmen t ( i . e . , a c hange i n p sycho logy d e sp i te a c on s tan t m o rpho logy ) t h en t h i s c ou ld b e ac on t r ibu to ry f a c to r t o t h e e me rgence o f af o od p roduc ing e conomy s ho r t ly a f te r t h e o n se t o f t h e p r e sen t i n t e rg lac ia l .

When D id t he P sycho soma t ic P henomena E vo lve? T he re a r et wo p e r iod si nw h ich p sycho soma t i c p henomena c ou ld h ave e vo lved : e i ther i n t h e f o od c o l lec t ing e nv i ronmen t o f t h e P a la eo l i th ic w he r e c o -ope ra t ion w a s b io log i ca l ly a dap t ive o r i nt h e f ood p roduc ing /u rban e nv i ronmen t w h i ch h a s g r adua l ly d eve lop ed s i n ce t h e N eo l i th ic a nd w he re r e g iona l c ompe t i t ion h a s b e en b i o log ica l ly a dap t ive . I ne i the r c a se , t h e p sycho log ica l s t a te s w h ich w ou ld b e i n d i ca t ive o f b i o log ica l u t i l i ty , a nd s oe vo lve t op romo t eo rh a rm p hy s ica lh ea l th , w ou ld b e r a the r d i f f icu l t . I f p sycho soma t ic p henomena e vo lved d u r ing t h e P a la eo l i th ic , t h en

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p sy cho log ica l s t a tes a s soc ia ted w i th c o -ope ra t ion , s uch a s f e e l ings o f s oc ia l s uppo r t , w ou ld b e h ea l thy a nd s t a te s a s soc ia ted w i th c ompe t i t ion , s uch a s a gg re ss ion t owa rd s o t he r s w ou ld b e u nhea l thy . B y c on t ra s t , i f p sycho soma t ic p henomena e vo lved i nf ood p roduc ing a nd u rban s o c ie t ies , t h en f e e l ings o f a gg re ss ion t owa rd s o t he r s a nd o the r m an if e s ta t ion s o f c ompe t i t ive s ucce s sw ou ld b e h ea l thy . I t i s m e r e ly n eces sa ry t o e xam ine t h e p sycho log ica l l i t e ra tu re o n t h e r e la t ionsh ip b e tween p sycho log ica l s t a tes a nd mo rb id i ty t os e e w he the r t h e d a ta f i t s b e t t e r t h e P a laeo l i th ic o r f ood p roduc ing /u rban mode l . T o s imp l ify t h e s omewha t c on t rove r s ia l r e su l t s , u nhea l thy p sy cho log ica l s t a tes i n c lude d ep res s ion ( J emmo t t & L ocke , 1 984 ) , p a r t icu la r ly s t a te s a s so c ia ted w i th b e reavemen t ( J ones , 1 987 ) , l o ne l ine ss ( Lynch , 1 977 ) , a nx ie ty ( F r iedman a nd B oo th K ew ley , 1 987 ) , h o s t i l i ty a nd a nge r ( Demb ro sk i a nd C o s ta , 1 987 ; F r iedman a nd B oo th -Kew ley , 1 987 ) , i n ab i l i ty t o e xp re ss e mo t ion i n c lud ing i n ab i l i ty t oe xp re s s h o s t i l i ty ( Sha f f e r e ta l , 1 987 ) , s u sp ic iousnes s ( Ba refoo t e t a l , 1 987 ) , a nd t i me p r es su re (W r igh t , 1 988 ) . H ea l thy p sycho log ica l s t a tes i n c lude s oc ia l s uppo r t ( Seeman a nd S yme , 1 987 ) , o p t im ism ( Sche ie r a nd C a rve r , 1 987 ) , a f f i l ia t ion mo t iva t ion ( J emmo t t , 1 987 ) a nd e f fe c t ive c op ing s t y les i n c lud ing h a rd ines s , c omm i tmen ta nd c on t ro l( Hu l l e ta l , 1 987 ) . I t w ou ld a ppea r t h a t a l though s ome o f t h e h ea l thy /unhea l thy p sycho log ica l s t a te s s uch a s d ep re s s ion a nd o p t im ism , f i t e qua l ly p sycho soma t ic e vo lu t ion d u r ing e i the r t h e P a laeo l i th ic o r t h e l a te r f ood p roduc ing /u rban s oc ie t ie s ,o the rp sycho log ica l s t a te s a r ec ons i s ten t o n ly w i th e vo lu t ion d u r ing t h e P a laeo l i th ic . C ons ide r , f o r e xamp le , t h e c ha rac te r is t ic o f h o s t i l i ty . H o s t i l i ty h as b een p r esen ted a s b e ing b i o log ica l ly a dap t ive ( Lo renz , 1 966 ) o n t h e b a s i s t h a t i tc on tr ibu tes t oi n d iv idua l i s t ic s e lec t ion , t h a t i s , s e le c t ion w he re o n ly t h e mo s t c ompe t i t ive m embe r s o f as pec ie s s u rv ive t o p a s s o n t h e i r g ene s . I nf ood p roduc ing a nd u rban e nv i ronmen t s w he re t h e re i sc ompe t i t ion f o r s ca r ce r e sou r ce s , t h e h o s t i le c ompe t i t ive i nd iv idua l s hou ld b e w e l l a dap ted -a s i lus t ra ted b y t h e s ucce s s o f " r obbe r b a ron s " . T hus , i f p sycho soma t ic p henomena e vo lved d u r ing f ood p roduc ing o r u rban e nv i ronmen t s , h o s t i l i ty s hou ld b e a n i n d ica to r o f b i o log ica l s ucces s a nd s hou ld b e h ea l thy -v e ry much i nt h e s ame w ay t h a t d om inance i sh ea l thy i n t h e r a inbow t r ou t ( Pe te r s e ta l , 1 988 ) . T ha t h o s t i l i ty i si nf a c t u nhea l thy i nh umans , a nd t h a t h o s t i l i ty w ou ld h ave b een m a ladap t ive i n a P a laeo l i th ic e nv i ronmen t a s i t w ou ld h ave r e duced c o -ope r a t ion , l e ad s t o t h e i n f e rence t h a t p sycho soma t ic p henomena e vo lved d u r ing t h e P a laeo l i th ic . S im i la r ly , o n ly P a laeo l i th ic e vo lu t ion i sc on s i s ten t w i th t h e f i nd ing t h a ts o c ia l s uppo r ta nd a f f i l ia t ion mo t iva t ion a r eh ea l thy . T hus t h e o ve ra l l p a t te rn o f t h e w ay p sycho log ica l s t a te s r e la te t o i l h ea l th i s

i n d ica t ive o f t h e e vo lu t ion o f p sycho soma t ic p henomena d u r ing t h e P a laeo l i th ic b u t n o t a t s ome l a te r p e r iod . T h i s i s b e cause a l l t h e se p sycho log ica l s t a te s c ou ld h ave e vo lved d u r ing t h eP a la eo l i th ic b u to n ly s ome c ou ld h ave e vo lved d u r ing l a t e r t i me s . T he c onc lus ion t o b e d rawn f r om t h e p sycho soma t ic d a ta i s a s f o l lows . P eop le b ecame i l d u r ing t h e t h e P a laeo l i th ic w hen t h ey e xpe r ienced c e r ta in s o r ts o f p sycho log ica l s t a te s o n a r egu la r b a s i s . P sycho soma t i c i n duced s i ckness d ur ing t h e P a laeo l i th ic w ou ld h ave b een a n a dap t ive m echan ism f o r e nsu r ing c o -ope ra t ion i n as o c ie ty w he re c o -op e ra t ion w a s e s sen t ia l f o r s u rv iva l . T he a dap t ive a dvan tage o f p sycho soma t ic i lne ss d i sappea r ed g r adua l ly w i th t h e d eve lopmen t o f f o od p roduc ing a nd u rban e nv i ronmen t s , t h ough t h e g ene t ic c apa c i ty t or e spond p sycho soma t ica l ly h a s n o t b een l o s t d u r ing t h e l a s t 8 ,000 y ea r s ( a r e la t ive ly s ho r t t i me p e r iod o f h uman e vo lu t ion ) . P eop le s t i l l r e a c t p sycho soma t ica l ly a s i ft h ey w e re i n a P a laeo l i th ic e nv i ronmen t . P sycho soma t i c i l ne s s i s l i ke a n a ppend ix : a s t ruc tu r e w h ich w a s o nce b i o log ica l ly a dap t ive b u tn ow a c t s s imp ly a s ah ea l th p rob lem .

T he U pper P a laeo l i th ic a nd t he E vo lu t ion o fL ongev i ty T he a r gumen t s o f a r h a s b een t h a t p sycho soma t ic p henomena e vo lved d u r ing t h e P a laeo l i th ic d ue t o t h e s t rong e vo lu t iona ry p r e s su r e t owa rd s c o -ope ra t ive b ehav iou r i nf ood c o l lec t ing s o c ie t ie s , p a r t icu la r ly u nde r c ond i t ions o f e nv i ronmen ta l s t res s . I n t h is s e c t ion w e s ugges t t h a t t h e g r ea tes t p r es su re t owa rds c o o pe ra t ion w ou ld h ave o ccu r red among s t U ppe r P a laeo l i th ic p eop le a nd t h e re fo re p sycho soma t ic p henomena a r e mo s t l i ke ly t o h ave e vo lved d u r ing t h eU ppe rP a laeo l i th ic . T he m ax imum l i f e e xp ec tancy o fa n o r gan ism i s g ene t i ca l ly c on t ro l led . ( The a ve rage l i f e e xpe c tancy i s ,o fc ou r se , c on t ro l led i na dd i t ion b y o t he r f a c to r s s u ch a s t h e p r e sence o f p r eda to r s ) . I n mo s t s pe c ie s t h e o nse t o f s ene scence i sl i nked t o ar e duc t ion i n r e p roduc t ive v a lue ( i . e . , t h e p robab i l i ty o f p roduc ing o f f sp r ing i n t h e f o l low ing y ea r ) . T he g ene t ic m e chan ism u nde r ly ing t h i s r e la t ionsh ip i st h a t g ene s c an b e " sw i tched o n " o n t h e b a s i s o f a n i n te rna l c l ock , i . e . ,t h eg ene h a s a n i mpac to n t h ep heno type o n ly w hen i ti ss w i tched o n . I fp a tho log ica l g ene s a r e s w i tched o n o n ly a f te rt h e o rgan ism h a s a ch ieved i t s f u l l r e p roduc t ive c apac i ty , t h en t h o se p a tho log i ca l g ene s a r en eve rs e lec ted o u t f r om t h e g ene p oo l . T hu s , o l d a ge i s r e f e r red t o a s a " gcne t ic d us tb in " b ecause o f t h e a ccumu la t ion o f p a tho log i ca l g ene s w h ich a r e e xp re s sed a t t h a t t i me ( P ianka , 1 988 ) . T he l o ngev i ty o f a na tom ica l ly m ode rn h uman s i s ab i o log ica l a noma ly . I ti sn o t t h a t t h e a v erage l i f e e xp ec tancy i st o o l o ng ( t he a ve rage l i f e e xpe c tancy i s , o f c ou r se , a f f ec ted b y t h e

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q ua l i ty o f med ica lc a re )b u tt h a t t h em ax imum l i f e e xpec tancy ( wh ich i sa bou t 1 00 ) o ccu r s w hen t h e r ep roduc t ive v a lue o f w omen h a s b e en z e ro f o r a bou t 6 0 y ea r s a nd t h e r ep roduc t ive v a lue o fm en h as b een l ow f o r al o ng p e r iod o f t i me . T he n o rma l e xp lana t ion ( e .g . B in fo rd , 1 983 ) f o r t h e e vo lu t ion o f l o ngev i ty i n a na tom ica l ly modem h umans i st h a t e l de r ly p eop le p l ayed s ome s o r to f s o c ia l r o le i n P a laeo l i th ic s o c ie ty , a s l e ade r s o r m ed ia to r s , a s e duca to r s o f t h e y oung o r a s r epo s i to r ies o f s oc ia l a nd e nv i ronmen ta l i n fo rma t ion . E v idence s ugge s t s t h a t t h e e vo lu t ion o f l o ngev i ty i s a r e cen t a nd r e la t ive ly a b rup t d eve lopmen t o f Homo s ap iens . U s ing a na ly s i s o f b one c hem i s t ry t o a s se s s t h e a ge o f d ea th i n N eander tha ls , T r inkaus a nd T homp son ( 1987 ) h ave s hown t h a t t h e o l de s t N eande r tha ls l i ved t o a t mo s t t h e e a r ly f o r t ie s - a max imum l i f e e xpec tancy w h ich , i n te re s t ing ly , i ss im i la r t ot h a t o f t h e c h impanzee ( Bu r ton , 1 962 ) . B y c on t r a s t , s ome a na tom ica l ly mode rn h uman s c on tempo raneous w i th t h o se N eande r tha l s a r e e s t ima ted t o h ave l i ved w e l l i n to t h e i r s i x th d ecade . T hus , t h e d eve lopmen t o f l o ngev i ty s e ems t ob e r e la t ive ly r e cen t a nd ac ha rac te r i s t ic o n ly o fH omo s ap iens s ap iens . I t i s u n l ike ly t h a t s u ch a n a noma lou s d eve lopmen t i n l o ngev i ty c ou ld h ave o ccu r red w i thou t g ood r e a son . A l though i ts e ems p robab le t h a t t h e re w e re f ew e l de r ly p eop le i nt h e U ppe r P a laeo l i th ic , t h e re w e re s ome a nd t h a t s ma l l n umbe r o f e l de r ly ( i . e . p o s t rep roduc t ive ) p eop le m us t h ave h ad s uf f ic ien t b io log ica l u t i l i ty t o c oun te r t h e n o rma l s enescence rep rodu c t ive v a lue r e la t ionsh ip i n e vo lu t ion . A s p hy s ica l s k i l l s d ec rea se a f te r f o r ty , t h e i d ea o f as o c ia l f u nc t ion o f e l de r ly p eop le s e ems mo s t l i ke ly , t h ough t h e p rec ise f unc t ion s uch i n d iv idua ls p l ayed i sj u s t s pecu la t ion . Wha t w e c an b e r e a sonab ly c e r ta in a bou t , h oweve r , i st h a t e l de r ly U ppe r P a la eo l i th ic p eop le c on t r ibu ted i ns ome w ay t ot h e s u rv iva l o f t h e g roup , a nd t h a t t h e i r c on t r ibu t ion w a s e s sen t ia l . A f te r a l l , d esp i te b e ing p hy s i ca l ly s t ronge r , a nd d esp i te h av ing a s im i la r ly s i zed b r a in , t h e N eande r tha l s d id n o t s ur v ive -b u t t h en t h ey d i dn o th ave e l de r ly p eop le . Wha teve r s oc ia l r o le e l de r ly p eop le h ad i n U ppe r P a laeo l i th ic g roup s , t h a t r o le c ou ld h ave b een u se fu l o n ly i ft h e re w a sc o -op e ra t ion . S o c ia l r o le s s uch a s l e ade r sh ip , m ed ia t ion , e duca t ion o r k now ledge r epo s i to r ie s c an b e f u lf i l led o n ly i f o the rm embe r s o f ag roup c o -ope ra te t oa l low t h e e l de r ly p e r son t o c a r ry o u t t h a t r o le . T he i n t roduc t ion o f e l de r ly p eop le i n t h e U ppe r P a laeo l i th ic w ou ld t h e re fo re h ave i n t roduced a n a dd i t iona l s t r a in t owa rd s c o -ope ra t ion s o i tw ou ld h ave b een a dd i t iona l ly a dvan tageous i f u n c oope ra t ive i n d iv idua ls b ecame i l a nd d i ed . Mo reove r , g i ven t h e p o s s ib le r o le s i d en t if ied f o r e l de r ly p eop le , i ti sc l ea r t h a t f ew e l de r ly p eop le w ou ld a c tua l ly b e n eeded , a nd t h a te xces s e l de r ly p eop le w ou ld h ave b een a d r a in o n r e sou r ce s .

T hu s , i tw ou ld a l so b e b i o log ica l ly a dap t ive i f e l de r ly p eop le c ou ld b e s e le c ted i nt e rms o f t h e i r c on t r ibu t ion t o t h e g roup s o t h a t g ood c on tr ibu to r ss u rv ive a nd o t he r sd on o t . I n s umma ry , d ur ing h uman e vo lu t ion , t h e U ppe r P a laeo l i th ic i s t h e p e r iod w hen p sycho soma t ic p henomena wou ld h ave b een mo s t u se fu l a s p romo te r s o f b io log ica l u t i l i ty . T he mo s t l i ke ly t ime f o r t h e e vo lu t ion o f t h e r e la t ionsh ip b e tween p sycho log ica l s t a te s a nd mo rb id i ty t h e r efo r e s e ems t o b e t h e Uppe r P a laeo l i th ic . A f te r t h e P a laeo l i th ic a nd Me so l i th ic , t h e b i o log ica l a dvan tage o f t h e p a r t icu la r p sycho log ica l s t a te -mo rb id i ty r e la t ionsh ip w h ich w e o b se rve h a s d ec r ea sed a nd a f te r t h eN eo l i th ic ad i f f e ren t f o rm o fr e la t ion sh ip m ay h ave b een a dap t ive . I ti sc er ta in ly p o ss ib le t h a tt h ep sycho log ica l s t a te -mo rb id i ty r e la t ion sh ip w e o b se rve i n h uman s t o day i s av e s t ig ia l a nd w eakened f o rm , a nd t h a t t h i s r e la t ionsh ip w a s s t ronge r i n t h e U ppe r P a laeo l i th ic . I t i s a l so p o s s ib le t h a t p sycho soma t ic p henomena a r e s t ronge r i n mode rn d ay f ood c o l lec t ing g roup s c ompa r ed t o g roup s w i th a f ood p roduc ing h i s to ry .

S ummary a nd P o s t scr ip t T he f unc t iona l t h eo ry o f i l ne s s s ugge s ts t h a t t h e r e la t ion sh ip b e tween p sycho log ica l s t a te s a nd o rp sy chosoma t ic p henomena e vo lved d u r ing t h e U ppe r P a laeo l i th ic a t w h ich t ime p r esenescen t i l ne s s a nd d ea th w e re b io log ica l ly a dap t ive . P sycho soma t ic p henomena w e re b i o log ica l ly a dap t ive ( i . e . e nhanced t h e s u rv iva l o f t h e g ene p oo l ) b y p ro tec t ing t h e s o c ia l c ohe s ivenes s o f g roup s o pe ra t ing i n a h a r sh e nv ironmen t . E v idence f o r t h i s t h eo ry c ome s f r om t h e p a r t icu la r p a t te rn o f p sycho log ica l s t a te s w h ich r e la te t o i lne s s , f r om i n f e rence s a bou t t h e e vo lu t iona ry f o r ces a f f e c t ing h uman e vo lu t ion d u r ing a nd s i nce t h e P a laeo l i th ic , a nd i n f e rence s a bou tt h ee vo lu t ion o fl o ngev i ty . T he f unc t iona l t h eo ry o f i lne s s d i f f e r s f r om s o c iob io log ica l t h eo r ie s w he re t r a i t s ( s uch a s c o o pe ra t ivenes s o r c ompe t i t ivenes s ) a r e a s sumed t o b e g ene t ica l ly d e te rm ined a nd b io log ica l ly a dvan tageous . We make n o a s sump t ion s a bou t t h e e x ten t t o w h ich t r a i t s a r e g ene t ica l ly o r e nv i ronmen ta l ly c on t ro l led . I n te re s t ing ly , i fs uch t r a i ts a re g ene t ica l ly c on t ro l led , t h en f unc t iona l p sycho soma t ic p henomena w ou ld h ave a r e la t ive ly s ma l l e f f ec t o n s uch t r a i t s i nt h e g ene p oo l a s s e lec t ion o f p eop le m ay o ccu r o nce t h ey h ave b ecome p o s t rep roduc t ive . H oweve r , t h e f unc t iona l t h eo ry o f i lne ss p rov ide s a n i d ea l a dap t ive m echan ism i f c o o pe ra t ion a nd h o s t i l i ty a r es i tua t iona l ly c on t ro l led ( i . e . , n o t g ene t ica l ly c on t ro l led , s e e L eakey a nd L ew in , 1 977 ) . S ucce s sfu l a dap ta t ion i na n U ppe r P a laeo l i th ic l i f e s t y le w ou ld h ave i n vo lved h o s t i l i ty t owa rd s o t he r a n ima l s b u t c o -ope ra t ion t owa rd s p eop le . T hu s , s ucce s s fu l a dap ta t ion

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D emb ro sk i , T .M . a nd C o s ta , P .T . ( 1987 ) . C o rona ry p r one b ehav io r : c omponen t s o f t h e T yp e Ap a t t er n a nd h o s t i l i ty . J .P e r s .5 5 :211-35 . F r iedman , H .S . a nd B oo th -Kew ley , S . ( 1 987 ) . T he " D i sea se -P rone P e r sona l i ty " : am e ta -ana ly t ic v i ew o f t h ec on s t ruc t . Am .P sycho ! .4 2 :539 -55 . G amb le , C . ( 1 986 ) . T he P a laeo l i th ic S e t t l emen t o f E u rope . C amb r idge : C amb r idge U n ive r s i ty P r e s s . G e i s t , V . ( 1 978 ) . L i f e S t ra teg ies , H uman E vo lu t ion , E nv i ronmen ta lD e s ign . N ew Y o rk : S p r inge rVe r lag . H a r r i s , G . a nd Mo rgan , M . ( 1 989 ) . S ucce s sf u l S ea T rou tA ng l ing . L ondon : B landf o rd . H u l l , J .G . , V an T reu r en , R .R . & V ir ne l l i , S . ( 1 987 ) . H a rd ine s s a nd h ea l th : ac r i t ique a nd a l te rna t ive a pp roach . J .P e rs .S oc .P sycho ! .5 3 :518 -30 . J emmo t t I I , J . B . ( 1 987 ) . S oc ia l mo t ive s a nd s u scep t ib i l i ty t o d i sea se : s t a lk ing i n d iv idua l d i f f e r ence si nh ea l th r i sk s . J .P e rs .5 5 :267 -98 . J emmo t tI l ,J . B . a nd L ocke , S .E . ( 1984 ) . P sycho soc ia l f a c to r s , i mmuno log ic m ed ia t ion a nd h uman s u scep t ib i l i ty t o i n f ec t iou s d i sea se s : h ow m uch d o w ek now? P s ycho ! .B u l l .9 5 :78 -108 . J one s , D .R . ( 1 987 ) . H ea r t d i sea se mo r ta l i ty f o l low ing w idowhood : s ome r e su l t s f r om t h e OPCS l o ng i tud ina ls t udy . J .P sycho som .R e s .3 1 :325 -33 . L eakey , R .E . a nd L ew in , R . ( 1 977 ) . O r ig ins . L ondon : Macdona ld a nd J ane ' s . L o renz , K . ( 1 966 ) . O n A ggre s s ion . L ondon : Me thuen . L ynch , J . J . ( 1 977 ) . T he B roken H ea r t : Med ica l C on sequence s o f L one l ine s s . N ew Y o rk : B a s ic B ook s . Me ik le john , C . , S chen tag , C .T . , V enema , A . &K ey , P . ( 1 984 ) . S oc ioeconom ic c hange a nd p a t te rn s o f p a tho logy a nd v a r ia t ion i n t h e Me so l i th ic a nd N eo l i th ic o f We s t er n E u rope : s ome s ugge s t ion s . I n P a leopa tho logy a t t h e O r ig in s o fA g r icu l tu re , e d s . M .N . C ohen a nd G .J . A rme lago s . p p . 7 5 100 . O r lando : A cadem ic P re s s . N ewe l l , R .R . ( 1 984 ) . O n t h e m e so l i th ic c on t r ibu t ion t o t h es o c ia le vo lu t ion o f We s te rn E u ropean S oc ie ty . I n E u ropean S oc ia l E vo lu t ion : A rchaeo log ica l P e r spec t ive s , e d . J .B in t l if f . p p . 6 9 -82 . B rad fo rd : U n ive r s i ty o fB radf o rd P r e s s . P e te r s , G . , F a i sa l , M . , L ang , T . & A hmed , I .( 1 988 ) . S t re s s c au sed b y s o c ia l i n te rac t ion a nd i t s e f f ec t o n s u scep t ib i l i ty t oA e romona s h yd roph i la i n f ec t ion i n r a inbow t r ou t S a lmo G a irdne r i . D i s . A qua t . O rg . 4 : 83 -9 . P i an l ca , E .R . ( 1 988 ) . E vo lu t ionary E co logy ( 3 rd e d . ) N ew Y o rk : H a rp e ra nd R ow . R enf r ew , A .C . ( 1 972 ) . T he Eme rgence o fC iv i l i za t ion : t h e C yc lad e s a nd t h e A egean i nt h e T h i rd M i l len ium BC . L ondon : Me thuen . R u sh ton , J . P . , R u s se l l , R . J .H . & We l l s , P .A . ( 1 984 ) . G ene t ic s im i la r i ty t h eo ry : b eyond k i n s e lec t ion . B ehay .G ene t . 1 4 :179 -93 . S ah l in s , M .D . ( 1 968 ) . T r ibesmen . L ondon : P ren t ice H a l l . S che ie r , M .F . a nd C a rve r , C .S . ( 1 987 ) . D i spo s i t iona l o p t im i sm a nd p hy s ica l w e l l -b e ing : t h e i nf luence o f g ene ra l ized o u t come e xpec tanc ie s o n h ea l th . J .P e r s . 5 5 :169 -210 . S e eman , T .E . a nd S yme , S .L . ( 1987 ) . S oc ia l n e two rk s a nd c o rona ry a r t e ry d i sea se : ac ompa r ison o f t h e s t ruc tu re a nd f u nc t ion o fs o c ia lr e la t ion s a sp r ed ic to r s o fd i sea se . P s ycho som . Med .4 9 :341-54 . S e rv ice , E .R . ( 1 966 ) . T he H un te rs . L ondon : P ren t ice H a l l .

r e qu i r e s s ome s i tua t ion s pec if ic h o s t i l i ty , t h a t i s , h o s t i l i ty w h ich o ccu r s o n ly i n a pp rop r ia t e s i t ua t ion s . A p e r son w ho w a s g ene t i ca l ly p r ed i spo sed t ob e a lway s n on -ho s t i le o r a lway s h o s t i le w ou ld b e b i o log ica l ly u nadap t ed t o t h e e nv i ronmen t . T he f u nc t iona l t h eo ry o f i l ne s s w ou ld p r ov ide am e chan i sm f o re l im ina t ing t h o se b i o log ica l ly u nadap ted i n d iv idua l s w ho w e r e i n app rop r ia t e ly h o s t i le , b u t w i thou t r e ly ing o n a c hange t ot h eg ene p oo l . T he f u nc t iona l t h eo ry o f i l ne s s p r e sen t s a v i ew w he re p r e sene s cen t d ea th a nd i l ne s s i s b i o log ica l ly a dap t ive . T o s ome p eop le t h is v i ew i sc oun te r in tu i t ive a nd t h i s i ss o b ecau se i ti s c ommon t o t a ke a p henomeno log ica l a nd i n d iv idua l i s t ic p e r sp ec t ive o f d ea th . E v idence f r om t h e s t udy o fo t he ra n ima l s s how s , h oweve r , t h a t p r e sene s cen t d ea th c an b e b i o log ica l ly a dap t ive . T ho se w ho p e r ce ive t h e f u n c t iona l t h eo ry o f i l ne s s t o b e af a nc if u l a nd i mp lau s ib le s t o ry s h ou ld c on s id e r t h e a l t e rna t ive . E i the r t h e p s y cho log i ca l s t a t e -mo rb id i ty r e la t ion sh ip e vo lved i nr e spon se t oe vo lu t iona ry p r e s su r e so ri t e vo lved b y c han ce . T o s ugge s t t h a t t h is r e la t ion sh ip e vo lved b y c hance g oe s a ga in s t t h e v e ry p r in c ip le o fs c ien t i f ic i n ve s t iga t ion . T o s a y t h a t s ome th ing o ccu r s " b y c hance " i ss c ien t i f ic n i h i l i sm . T h ing s w h ich a r e t h ough t t oo ccu r b y c hance a r e o f ten s hown a t al a t e r d a t e t o o bey n o rma l s c ien t if i c l aw s . T he " i t o c cu r r ed b y c hance " e xp lana t ion i ss imp ly n o t a n e xp lana t ion f o r t h e e vo lu t ion o f p sycho soma t i c p henomena . T he f u nc t iona l t h eo ry o f i l ne s s m ay n o t b e c o r r ec t ; b u t i ti st h e o n ly t h eo ry o ft h e e vo lu t ion o fp sycho soma t ic p henomena w h ich e x i s t st o day .

R e f erence s B a i ley , K . ( 1 987 ) . H uman P a l eop sycho logy : A pp l ica t ion s t o A ggres s ion a nd P a tho log ica l P roce s ses . S an F ran s i sco : E r lbaum . B ake r , G .H .B . ( 1 987 ) . P sycho log ica l f a c to r s a nd i mmun i ty . J .P sycho som .R es .3 1 :1-10 . B a re foo t , J . C . , S i eg le r , I . C . , N ow l in , J . B . , P e t e r son , B .L . , H aney , T .L . , & W i l l iams J r . , J . B . ( 1 987 ) . S u sp ic iou sne s s , h ea l th a nd m o r ta l i ty ; af o l low -up s t udy o f5 00 o l de r a du l t s . P s ycho som . Med . 4 9 :450 5 7 . B a te son , G . ( 1 979 ) . M ind a nd N a ture . N ew Y o rk : D u t ton . B ende r , B . ( 1 975 ) . F a rm ing i n H is tor y : F rom H un te r /Ga the re r t oF ood P roducer . L ondon : J ohn B ake r . B in f o rd , L .R . ( 1 983 ) . I nP u rsu i t o ft h e P a s t . L ondon : T hame s a nd H ud son . B u r ton , M . ( 1 962 ) . U n ive rs i ty D ic t iona ry o f Mamma l s o ft h e Wo r ld . N ew Y o rk : C romwe l l . C h r i s t ian , J . J . ( 1980 ) . E ndoc r ine f a c to r s i np opu la t ion r e gu la t ion . I n B io soc ia l Mechan i sm s o fP opu la t ion R egu la t ion . e d s . M .N . C ohen , R .S . Ma lpa s &H .G . K le in . p p .55 -115 . Y a le : Y a le U n ive r s i ty P r e s s . C ou s t eau , J .Y . ( 1975 ) . T he A c t ion o fL if e . N ew Y o rk : P l enum .

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S ha f f e r , J .W . , G rave s , P .L . , S wank , R .T . & P ea r son , T .A . ( 1987 ) . C lu s te r ing o fp e r sona l i ty t r a i t s i ny ou th a nd t h e s ub sequen t d eve lopmen t o f c ance r a mong p hy s ic ians . J .B ehay . Med . / 0 :44 1 -47 . S t r inge r , C .B . ( 1988 ) . T he d a t e s o f E den . N a ture 3 3/ :565 -66 . S t r inge r , C .B . , G run , R . , S chwa rcz , H .P . &G o ldb e rg , P .( 1989 ) . E SR d a te sf o rt h eh om in id b u r ia ls i te o fE s S khu l i nI s rae l . N a tu re 3 38 :576 -78 .

T r inkau s , E . a nd T homp son , D .D . ( 1987 ) . F emo ra l d i aphy sea l h i s tomo rphome t r ic a ge d e te rm ina t ion s f o r t h eS han ida r3 ,4 , 5a nd 6N eande r ta l s a nd N eande r ta l l o ngev i ty . Am .J .P hys .A n thropo l . 7 2 :123 -29 . Wh i te , L .A . ( 1 959 ) . T he E vo lu t ion o fC u l ture . N ew Y o rk : McG raw -H i l l . Wr igh t , L . ( 1 988 ) . T he T ype A b ehav io r p a t te rn a nd c o rona ry a r te ry d i sea se . Am .P sycho l .4 3 :2 -14 .

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4 .

HEALTH , ADAPTAT ION , AND MALADAPTAT ION I N PAST SOCIETIES

A lan H . G oodman r e cen t ly r e v i sed ( Goodman a nd A r rne lago s , 1 989 ) ( F igu r e4 . 2 ) . T he p u rpo se o ft h i s mode l i sn o t t o c od i fy l i nk s b e twe en c omponen t i nt h e a dap t ive p ro ce s s , o rt os ugge s t t h a t a l l a dap t ive d ynam ic s c an b e t r a ced t h rough t h e mode l . R a the r , t h e p u rpo se o f t h e m ode l i st op rov ide am echan i sm f o rs y s tema t ica l ly e xam in ing a nd r ef lec t ing o nt h e a dap t ive p ro ce s s a nd t h e c en t ra l i ty o f h ea l th t o t h i sp ro ce s s . T he m ode l s t a r t s w i th a c ons ide ra t ion o f " e nv i ronmen ta l c on s t ra in t s " t oa dap ta t ion ( F igu re 4 . 2 ) . T he se a r e t y p i ca l ly d i v ided b e tween l i m i t ing r e sou r ce s a nd s t r e s so r s . T he mo s t i mpo r tan t l i m i t ing r e sou r ce sa r e l i ke ly t ob e b a s ic o ne s : f o od , w a te r , s h e l te r . S t r e s so r s m ay b e r e la ted t oc l ima t ic e x t r eme s , s u ch a s e xce s se s o f h ea t a nd c o ld , o r l ow p a r t ia l p r e s su r e o f o xygen . A l l o f t h e se c l ima t ic a nd p hy s ica l s t re s so r s a r e i mpo r tan t b ecau se t h ey m ay h ave a n e f f ec t o n r e s i s tance t o d i sea se . O the r e nv i ronmen ta l s t re s so r s , s u ch a s p a r a s i te s a nd p r eda to r s , h ave m o re o bv iou s a nd d i r e c t a f f e c t s o n h ea l th a nd l o ngev i ty . T he r e a r e an umbe r o f d i f f e r en t w ay s t h a t e nv i ronmen ta l ly i n i t ia t ed c on s t ra in t s a nd s t r es so r c an b e a dap ted t o . C u l tu re , t h rough i t s t e chno log i ca l , s o c ia l a nd i d eo log ica l s y s tems , m ed ia te s t h ep ro ce s so fe x t r ac t ing r e sou r ce s f r om t h e e nv i ronmen t . C u l tu r a l s y s tems a r e g ene ra l ly e f f e c t ive i n b u f f e r ing e nv i ronmen ta l c ons t ra in ts a nd s t r e s so r s , o r r emov ing t h e i n d iv idua ls i n a s o c ie ty f r om c on tac tw i th t h es t re s so r . T he r e a r e n ume rou s e xamp le s o f e f f ec t ive c u l tu ra l b u f f e r ing o f s t r e s so r s a nd l i m i t ing r e sou r ce s . S he l te r a nd c l o th ing a r e g ene ra l ly w e l l s u i ted a s b u f f e r s a ga in s t e x t r eme s o f c o ld a nd h ea t . A n i n te r e s t ing e xamp le o f h ow s he l te r c an a i d a dap ta t ion i sp r e sen ted b y May ( 1 960 ) w ho r e la te s h ow p ea san t s i n t h e h i l ly r e g ion o f N o r th V ie tnam h ave c u s toma r i ly b u i l t t h e i r h ou se s o n s t i l t s , a bou t e i gh t t ot e n f e e t a bove t h e g r ound . T h i s i sj u s t a bove t h e f l igh t c e i l ing o f A nophe le s m in imu s , av e ry f i e r ce m a la r ia v e c to r . T he a dap t ive s i gn i f icance o f b u i ld ing h ou ses o n s t i l t s b e came e ven c l ea r e r w hen o ve rpopu la t ion d eve lop ed i n t h e mo re f e r t i le d e l ta r e g ion s a nd g r oup s f r om t h e d e l ta b egan t o m ig ra te t o t h e h i l l s . T he d e l ta p eop le s w e re a c cu s tomed t o b u i ld ing t h e i r h ou se s o n t h e g r ound . S ub sequen t ly , t h ey f o und t h em se lve s t ob e w i th in r a nge o f t h e m a la r ia v ec to r a nd t h ey s u f f e r ed f r equen t ly f r om m a la r ia . C on sequen t ly , t h e d e l ta p eop le s h ave b een d i s cou raged f r om r e lo ca t ing a nd a t t r ibu te t h e i r h ea l th p rob lems t o t h e e v i l s p i r i t si nt h eh i l l s( May , 1 960 ) .

I n troduc t ion T he a na ly s i so fh ea l th i np a s ts o c ie t ie s h a st o o f r equen t ly b een u nd e r taken i n av acuum . Many p a leopa tho log ica l s t ud ie s h ave b een u nde r taken w i th l i t t le c oncer n f o re i the rt h ec au se o re f f e c to f h ea l th p rob lem s . T oo i n f r equen t ly h ave e f fo r t s b een m ade t o a t temp t t o u nde r s t and t h e r e la t ionsh ip b e twe en h ea l t h a nd a dap ta t ion , a nd e qua l ly i n f requen ta r ea na ly se so ft h ec au se so fi l h ea l th t h a t f o cu s b eyond t h e i mmed ia te e nv i ronmen t . P a leopa tho log i s t s a r e g u i l ty o f " q u i t t ing e a r ly " ( Ha r r is , 1 988 ) , b e fo r e t h e i mp l ica t ion s o f s i gns o f h ea l th a nd i l ne s s h ave b een f u l ly u nde r s tood . P a leopa tho log i ca la na ly se s , n eve r the le s s , h ave g r ea t p o ten t ia l f o r e l uc ida t ing p a s t a dap t ive s t rugg le s , a nd t h e i r l i nk s t o t h e p r e sen t h uma n c ond i t ion . I n d iv idua l m ea su re s o fh ea l th d e r ived f r om s ke le ta l r ema in s , r a ng ing f r om m i ld g row th r e ta rda t ion t om o r ta l i ty ,c an b ev i ewed a ss i gn so f t h e s t rugg le t oa dap t . T he p a t ter n o f i n d iv idua l i n d ica t ion s o fh ea l th a nd d i sea se p rov ide s ab a s i s f r om w h ich o ne c an i n f e r t h e a dap t ive a nd f u nc t iona l c o s t s o f i l ne s s i n a r ea s s u ch a s r e p roduc t ion , r e s i s tance t o d i sea se , s o c iab i l i ty , w o rk c apa c i ty , a nd l e a rn ing . T he p a t te rn o f d i sea se i n g roup s c an b e u s ed t o r e con s t ru c t d i f f e ren t ia l a c ce s s t o r e sou r ce s a nd e xpo su r e t o s t r e s s . I nt h e f o l low ing p ap e r Iw i sh t op rov ide s ome s e n se o f t h e p o t en t ia l o f p a laeopa tho logy , a nd s ome s en se o ft h e c ha l lenge o fn o t q u i t t ing e a r ly . Ib eg in b y i n t roduc ing a mod e l f o r s t udy ing h ea l th a nd d i sea se i nr e la t ion sh ip t oa dap ta t ion i n p a s t s oc ie t ie s . A f te r t h e m ode l i s b r ief ly e xp la ined , t h e f u nc t iona l s i gn i f i cance o f g row th s t un t ing , o ne m ea su r e o f h ea l th s t a tu s , i s e xp lo red . T h is d i s cu s s ion i sb a sed o n s t ud ie s o f t h e f u nc t iona l c on sequence s o f s ho r t s t a tu r e i n c on t empo ra ry p opu la t ion s a nd i s m ean t t o i l u s t ra te t h e t y p e o f i n f e r ence s t h a t c an b e d e r ived f r om m ea su r emen t so fb one s a nd t e e th . I e nd b y s ugge s t ing d i r e c t ion s f o r f u tu re r e sea r ch a nd b y p l a c ing t h e s t udy o fh ea l th a nd a dap ta t ion i nt h e l a rge rt h eo r e t ica ld eba t eo ve rt h ev a l id i ty o f p r o ce s sua la pp roa che si na r chaeo logy .

A Mode l o fH ea l th , Nu tr i t ion ,a nd A dap ta t ion i nP as tS co c ie t ie s T h i sa dap t iv i s tm ode l w a so r ig ina l ly p r e sen ted i n1 982 ( Hu ss -A shmo re e ta l . , 1 982 )a nd h a sb een

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A s a g ene r a l e xamp le , t h e a dop t ion o f a g r icu l tu re i s a n e conom ic a d ju s tmen t t h a t e mbod ie s an umbe ro fc hange s i nt h e a b i l i ty o fa c u l tu r e t o b uf f e r s t r e s se s . T he e ne rge t ic e f f i c iency o fa g r i cu l tu re a nd t h e a moun t o ff o od p r oduced p e r u n i t a r ea i s g r ea te r t h an t h a t o f h un t ing a nd g a the r ing . T hu s , o n t h e f a ce o f i t , a g r icu l tu r e c an p rov ide a b u f f e r a ga in s t m a lnu t r i t ion , p e rhap s t h e m o s t p e r s i s t en t a nd s i gn if i can t s t r e s so r f a ced b y p a leo l i th ic h un te r g a the re r s ( McKeown , 1 988 ) . H oweve r , o ne n ea r ly i n va r iab le d emog raph i c e f f e c t o f a g r icu l tu ra l i n t en s i f ica t ion i sg r ea te r p opu la t ion d ens i ty . U nfo r tuna te ly , t h e i n c r ea sed p opu la t ion d en s i ty a nd o t he r e co log i ca l a nd d emog r aph i c c hange s t h a t a r e o f ten s e conda ry t oa g r icu l tu ra l i n ten s if ica t ion h ad u nexpec ted c on sequence s f o r h ea l th , s u ch a s i mp rov ing c ond i t ion s f o r t h e s p r ead o f i n f ec t iou s d i sea se s . F u r the rmo r e , i n We s t A f r ica i ti sl i ke ly t h a t c l ea r ing o ff o r e s t f o r a g r i cu l tu ra lp r oduc t ion b r ough th uman si n to c l o se c on ta c t w i th m o squ i toe s , t h u s l e ad ing t o t h e s p read o fm a la r ia i nt h a ta r ea ( L iv ing s tone , 1 9 58 ) . T he i mpo r tan t p o in t i nt h e a bove i st h a t t h e d ynam ic c u l tu ra ls y s tem ,w h ich Ih ave j u s tb r ief ly n o ted , o pe ra te s i n t wo w ay s : a s ab uf f e r ing s y s t em a nd a s ap roduce r o f n ew s t r e s so r s a nd c ons t r a in t s . A s c u l tu re s c hange t h e n a tu r e o f e nv i ronmen ta l a dap ta t ion c hange s . I mp roved a d ju s tmen t t o o ne e nv i ronmen ta l c ond i t ion i s f r equen t ly o n ly m ade b y i n c r ea s ing e xpo su r e t o a no the r c ond i t ion . C omp rom i sed a dap ta t ion s a r e f r equen t ; t h ey a r e t h e r u le . A s w e l l , c u l tu r e sa nd t h e i r a dap ta t ion s v a ry i n h ow w e l l d i f f e r en t s e gmen t s o f t h e p opu la t ion a r e b u f f e r ed f r om s t re s so r s( Goodman a nd A rme lago s , 1 989 ) . T he e nv i ronmen ta l c on s t ra in t s t h a t a r e n o t w e l l b uf f e r ed b y t h e c u l tu ra l s y s tem , a l ong w i th t h e n ew ly p roduced c u l tu ra l s t r e s so r s , r e a ch a nd c on ta c t t h e i n d iv idua ls i nt h e s o c ie ty . F rom t h i s p o in to n ,t h ed eg ree t ow h ich t h e se c on s t ra in t sa r e a dap ted t o d epend s o n t h e i n d iv idua l ' s l e ve l o f h o s tr e s i s tance . R e s i s tance c an b er e la t ed t oo ne ' s g ene t ic , d eve lopmen ta l , a nd p hy s io log ica l s t a tu s . I f as t re s so ri sp e r s i s t en ta nd l i f e t h r ea ten ing o ve r m i l lenn ia t h en ag ene t ic r e s i s tance m igh td eve lop . T h i s a ppea r s t o h ave b e en t h e c a se a lmo s t u n ique ly i n We s t A f r ica w he r e t h e f r equency o f t h e Sh emog lob in a le le ,w h ich l e ad st os i ck le c e l l t r a i t , i n c r ea sed g ene t i c r e s i s tance t o m a la r ia ( L iv ings tone , 1 9 58 ) . Mo re o f t en , h oweve r , s t re s se s a r e n o t s o p e r s i s t en t a nd a g ene t ic " s o lu t ion " i s n eve r d eve loped . I n t h e se c a se s s ome d eg r ee o f r e s i s tance a nd a dap ta t ion m ay d e r ive f r om s ho r t e r t e rm d eve lopmen ta l o r p hy s io log ica lc hange s( Thoma se ta l . , 1 9 79 ) . U nfo r tuna te ly , c h ron ic s t r e s so r s a nd l i m i t ing r e sou r ce s o f ten o ve rbu rden d eve lopmen ta l a nd p hy s io log ica l s y s tems . T hu s , w h i le s u ch s y s tem s a r e i n i t ia l ly a b le t o m ee t e nv i ronmen ta l c ha l lenge s , t h ep e r s i s tence o ft h ec ond i t ion so f ten r e su l t si n ad ec rea sed h ea l th a nd n u t r i t iona ls t a tu s , w h i ch w i l lb ef u r the rl owe r ed b ye xpo su r et on ew

s t r e s so r s . I n d iv idua l s w ho h ave p oo r n u t r i t ion , f o r e xamp le , a r e l e s s r e s i s tan t t o i n f ec t iou s d i sea se s , a nd i n f e c t iou s d i sea se f u r the r l owe r s n u t r i t iona l s t a tu s ( Ma r to r e l l , 1 980 ; A l len , 1 9 84 ; Ma ta e ta l . , 1 9 71 ) . T he s e v e r i ty a nd d u ra t ion o f t h e s t r e s s r e spon se m ay b e v i ewed a s af u nc t ion o f t h e d eg r ee o fc u l tu ra l a nd e nv i ronmen ta l c on s t ra in t s a nd s t re s so r s , b a lanced a ga in s t t h e a dequa cy o f t h e c u l tu ra l b u f f e r ing s y s tem a nd i n d iv idua l r e s i s tance r e sou r ce s . F o r tuna te ly f o r t h e p a leopa tho log i s t , t h e s t r e s s r e spon se , s t e ro typ ic p hy s io log ica lc hange s r e su l t ing f r om t h e s t rugg le t o a d ju s t , i s f r equen t ly m an i f e s t i n r e la t ive ly p e rmanen t s k e le ta l c hange s . Wh i le s ome s t r e s so r s , i fr a p id a nd s y s tem s pec i f ic , m ay n o t l e ave a ny e v idence o f as k e le ta lr e spon se ( O r tne r , 1 9 90 ) , m any s t r e s so r d o l e ave s c a r s o n s k e le ta l e l emen t s . B e cau se b one a nd t e e th a r e l i m i t ed i n t h e i r r e spon s e r e pe r to i r e , o ne c an r e ad t h a t a s t r ugg le t oa d ap t h a s o ccu r red , a l though l e s s i n k nown o f t h e s t r e s so r t h a t w a s t h e c a u s e o f t h e s t rugg le . H ea l th a nd a dap ta t ion h ave s i gn i f icance t h a t e x tend s b eyond t h e i n d iv idua l s t ot h e p opu la t ion a nd s o c ie ty . A s w e h ave o u t l ined i nt h e m ode l , s t r e s se s e f f e c t p opu la t ion s a nd g r oup s b y d e c r ea s ing w o rk c apa c i ty , d ec rea s ing f e r t i l i ty , i n c r ea s ing m o rb id i ty a nd m o r ta l i ty , a nd c a us ing s e conda ry d i s rup t ion s t ot h e s o c ia l , p o l i t i ca l , a nd e conom i cs t r uc tu r eo f ac ommun i ty ( A l len , 1 984 ) . I n t u rn , t h e se c u l tu ra l a nd p opu la t io n l e ve l c hange sc y c le b a ck t oc au se f u r the rc hange si nt h e e nv i ronmen ta nd c u l tu ra ls y s tem . T he e vo lu t ion o ft h i s m od e l f r om t h ee a r ly t o l a te 1 980 sr e f l e c t s a ne xpan s ion i nt h ink ing a bou t t h e r o le o f h e a l th a nd i t s s i gn i f icance i nh uman a dap ta t ion . I nt h ee a r l ie rm ode l , t h ee nd r e su l to f af a i led s t r ugg le t oa dap t w a s t h e d ea th o f t h e i n d iv idua l ( Hu s s -A shmo r e e ta l . , 1 9 82 ) . I nt h e r e v i sed m od e l a f u r the r s t ep w a s a dded t o d emon s t r a te h ow t h ei n d iv idua l ly b a sed i n d i ca to r s o f s t r e s s , t h e p a leopa tho log i ca l d a ta , m igh t b e u s ed t om ak e i n f e r ence s a tt h e p opu la t ion l e ve l ( Goodman a nd A rme lago s , 1 989 ) . A s w e l l , t h e f e edba ck a r row s w e r e a dded t oi l u s t r a te h ow t h e s t r e s so r s c ou ld h ave a f u r the r c yc l ica l e f f ec t o n e nv i ronmen t a la nd c u l tu ra ls y s tems . I ti s ac h a l lenge t o p a leopa tho log i s t s t o g o b eyond t h ei n d iv idua lt oe xam ine h ow p a t te rns o f d i sea se a nd d i s rup t ion h ave s i gn if ica nc e f o r f am i l ie s , l a rge r s o c ia l g r oup s , a nd c u l tu r e s . S im i la r ly , i ti sac ha l lenge t og ob eyond c on ce rn s w i th p r ox ima t e c au se s o f i l ne s s ( c on ta c t w i th i n f ec t iou s p a thogen s , i n su f f ic ien t d i e t s ) t o e xam ina t ion o ft h eb r oade ra nd u nde r ly ing s o c ia l , e conom i c , a nd p o l i t ica l f a c to r s w h ich d e te rm ine w ho c ome si n to c on ta c tw i th s t re sso r sa nd w ho se r e sou r ce s a r e m o s t l i m i t ed . I ti sp ropo sed t h a t i t i s o n ly i n t h i s w ay t h a t p a leopa tho logy w i l l p r ov ide i n s igh t s i n to a dap ta t ion i nt h e p a s t . I ti s o n ly a s up e r f i c ia l p a radox , h oweve r , t h a t t h e i n f e r ence s u pon w h ich t h e se i n s igh t s n e ed t ob e

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t h a t t h e re a r e f ew f u nc t iona l i n f e r ence s t h a t c an b e m ade b a sed o n a n th ropome t r ic d a ta a nd w e n eed n o t w o r ry s o m uch a bou t t h e m i ld to mode ra te ly m a lnou r ished . C on t ra ry t o t h e " sma l l b u t h ea l thy " h ypo the s i s , Ma r to r e l l , i n a n e a r ly r e v iew o f t h e l i t e ra tur e o n g row th , n u t r i t iona l s t a tu s a nd i n f e c t iou s d i sea se , f o und t h a t p oo r g row th s t a tu s i s a lmo s t i n va r iab ly a s so c ia ted w i th i n c r ea sed d i sea se i nd eve lop ing c oun t r ie s ( Ma r to re l l , 1 980 ; a l so s e e T ompk in s , 1 988 ) . A l though t h i s r e la t ion sh ip a pp ea r s t ob es omewha ta t tenua ted i n d eve lop ed c oun t r ie s , t h e p i c tu r e o fn ea r ly a l l p a s t s o c ie t ie s i s m o re a k in t o t h e s i tua t ion i n c on tempo r a ry d eve lop ing t h an d eve loped c oun t r ie s . T he r e la t ion sh ip b e tween u nder nu t r i t ion a nd d i sea se i n d eve lop ing c oun t r ie s i ss o s t rong t h a t i ti sd i f f icu l t i n many c a se s t oa s ce r ta in t h e c au se o fp oo r g r ow th s t a tu s ( Ma r to r e l l , 1 989 ) . G row th s t a tu s s e ems t o b e a s i gn o f t h e c umu la t ive s t r e s s e xpe r ienced b y a n i n d iv idua l , o f w h ich p oo r d i e t a nd d i sea se a r e l i ke ly t ob e o fm a in i mpo r tance . Ar e v iew o ft h e l i t e ra tu r e c l ea r ly s how s t h a t t h e p r ima ry p rox ima te c au se s o f s ho r t s t a tu r e a r e u nde rnu t r i t ion a nd d i sea se . T o t h en a f f i rm t h a t s ho r t s t a tu r e i sa dap t ive i mp l ie s as a n c t ion ing o f t h e p r e sence o f t h e c ond i t ion s l e ad ing t o s ho r t s t a tu r e ( Ma r to r e l l , 1 989 ) , w h ich m igh t u l t ima te ly b e t r aced b a ck t op ove r ty a nd i n equa l i ty i na cce s s t or e sou r ce s( M cKeown , 1 988 ) . I n a dd i t ion t o i n c r ea sed m o rb id i ty , p oo r g r ow th s t a tu si sa s so c ia ted w i th an umbe ro fo t he r f u nc t iona l c on sequence s . C havez a nd Ma r t inez ( 1 982 ) h ave s y s tema t ica l ly a dd re s sed t h e f u nc t iona l c o s t o f u nde rnu t r i t ion a nd s ma l l b ody s i ze i n c h i ld r en f r om T ezon teopan , Mex ico . T he i rl o ng t e rm n u t r i t iona l s upp lemen ta t ion s t udy b egan i nt h e 1 960 s . P r egnan t m o the r s a nd t h e i r i n f an t sw e re d i v ided i n to t wo g r oup so fw h ich o ne r e ce ived d a i ly n u t r i t iona ls upp lemen ta t ion a nd t h e o t he rg r oup d i dn o t . T he se a u tho r s f o und t h a t t h e b e t te r n ou r i shed c h i ld r en c r y l e s s a nd t a lk a ta n e a r l ie r a ge . T he a moun t o f a c t iv i ty i s h i gh ly r e duced i nt h e s ho r te r c h i ld r en . T he s ho r te r a nd n on supp lemen ted c h i ld r en d i sp layed l e s s e xp lo r a to ry b ehav iou r ( Chavez a nd Ma r t inez , 1 982 ) . T h i s s t udy s how s t h a t g r ow th c ompe t e s w i th a c t iv i ty a l i m i ted n u t r ien t s upp ly . When g r ow th i s c omp rom i sed a c t iv i ty i s n o t s pa r ed . R a the r ,t h er e ve r se s e ems t ob e t r ue . P oo rg r ow th s ugge s t s t h a t a c t iv i ty h a s a l so b een a f f ec ted . T he r e a r e f u nc t iona l c o s t i n t e rms o f d ec rea sed a c t iv i ty a nd l e a rn ing t h a ta r e s i gna l led b y g row th s t un t ing ( F igu r e4 . 1 ) . I ti sa l so c l ea r f r om an umbe r o f s t ud ie s t h a t p oo r g row th i sr e la ted t o a n i n c rea sed r i sk o f m o r ta l i ty , t h e u l t ima te i n d ica to r o f a dap t ive f a i lu r e ( s e e A lam e ta l , 1 989 ; C hen e ta l . , 1 980 ; L e rbe rghe , 1 988 ; Smedman e ta l , 1 987 ) . A lam a nd c o -wo rke r s ( 1 989 ) f o re xamp le , h av er e cen t ly f o und t h a t c h i ld r en i n t h e a r ea o f T ekna f , B ang lad e sh w i th s ma l l a rm c i r cumf e r ence s s u f f e r

b a sed n eed t oc ome a tl e a s ti np a r tf r om s t ud ie so f c on tempo ra ry g r oup s .

N u tr i t ion , D isease ,a nd F unc t ion : T he S ign if icance o f Sma l l B ody S i ze S imp le a nd u ncomp l i ca ted a n th ropome t r ic m ea su remen t s a r e ap owe r f u l m ean s f o ra s se s s ing a p opu la t ion ' s a dap ta t ion t o e nv i ronmen ta l c ond i t ions . E xcep t f o r av e ry l i m i t ed n umbe r o f e t hn ic g roup s s u ch a s N i lo t ic A f r ican s , t h e r e i s s u rp r is ing ly l i t t le v a r ia t ion i nm ean h e igh ta mong d i f f e ren t e t hn i c g r oup s t h a t c an b e a t t r ibu ted t o g ene t ic d i f f e r ence s ( D ie tz , 1 983 ; G ra i t ce r a nd G en t ry , 1 981 ; H ab ich t e ta l . , 1 974 ) . T hu s , i ti s n ow a ccep t ed p rocedu r ef o rn u t r i t iona ls c ien t i s tt o c ompa re h e igh t s i n d i f f e r en t g roup s a ga in s t a u n ive r sa l s t anda rd ( Hab i ch t e t a l . , 1 974 ) . B ecau se o ft h e u n ive r sa l i ty o ft h e g r ow th p a t t er n a tt h e g roup l e ve l , i ti sp o s s ib le t oc ompa r e t h e p r eva lence o f s t un t ing i n g r oup s , d e f ined i n v a r ious w ay s s u ch a s t h e p e r cen t o f i n d iv idua l s b e low t h e 5 t h o r 1 0 th p e r cen t i le , b e low n ega t ive t wo s t anda rd d ev ia t ion s ( n ea rt h e 2 nd p e r cen t i le ) , o r b e low 8 0% o r 9 0% o f t h e m ean o f t h e s t anda rd . O ne s hou ld , f o re xamp le ,e xp ec ta bou t 5 % o fi n d iv idua l s t ob e b e low t h e f i f th p e r cen t i le h e igh t fo r -age d ue t or a ndom p ro ce s se s . I ft h e o b se rved p e r cen t i ss i gn if i can t ly g r ea t e r t h an 5 % t h en o ne c an u sua l ly i n f e r a n e nv i ronmen ta l c au se . I n t h i s w ay , t h e p a t te rn o f s t r e s s a mong a nd w i th in g roup s c an b e r e ad i ly c ompa r ed . I n t h e a na lys i s o f t h e p r eva lence o f s t un t ing , n u t r i t ion i s t s a nd i n te rna t iona l h ea l th r e sea r che r s h ave ap owe r fu l t o o l f o r t a rge t ing p opu la t ion s a t r i sk . Wh i le t h e g ene ra l p ro cedu r e f o r e va lua t ing s t un t ing h a s n eve r b een s e r iou s ly c ha l lenged , i n 1 980 t h e e conom i s t D av id S ek le r c ha l lenged a n umbe r o f f u ndamen ta l a s sump t ion s a bou t t h e m ean ing a nd c on sequence s o f s ho r t s t a tu r e i n d eve lop ing c oun t r ie s . S ek le r p ropo sed t h a t m o s t i n d iv idua l s t h a t a r e s ho r t i ns t a tu r e i nd eve lop ing c oun t r ie s a r e s ho r t b ecau se t h ey h ave a dap ted t o m a rg ina l f ood a va i lab i l i ty ( S ek le r , 1 980 , 1 982 ) . S ek le r a s se r ted t h a t t h e b ody a d ju s t s t o c h ron ica l ly l ow n u t r ien t a va i lab i l i ty b y r e du c ing i t s g r ow th . I n t h i s w ay i ti s a b le t o r e ga in p hy s io log ica l e qu i l ib r ium . S ek le r f u r the r s u gge s t s t h a t t h i s r e ad ju s tmen t o ccu r s w i thou t a dap t ive c o s t . T he se i n d iv idua l s a r e e s sen t ia l ly " sma l l b u t h ea l thy " ( S ek le r , 1 980 :225 ) . I n f a c t , s ma l l i n d iv idua l s m ay b e b e t te r a dap ted t h an l a rge r i n d iv idua l s t o a n e nv i ronmen t i n w h ich f o od s c a rc i ty i sac h ron ic p r ob lem . S ek le r ' s " sma l l b u t h ea l thy " h ypo the s i s h a s a t r a c t ed ag r ea t d ea l o fa t ten t ion a nd e ngend e red m u ch d eba te a mong p o l i cy m ake r s , n u t r i t ion i s t s , a n th ropo log i s t s a nd o t he r s c on ce rned w i th t h e i n t e rna t iona l d imen s ion o f m a lnu t r i t ion . T he m a in q ue s t ion c once rn s w he the r o r n o t t h e se i n d iv idua l s a r e , i n de ed , s ma l l b u t h ea l thy . I f S ek le ri sc o r r e c t i nh i sp ropo s i t ion t h en i tf o l low s

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a t we lve -fo ld i n c rea sed p robab i l i ty o f d y ing c ompa r ed t o c h i ld r en w i th m o re a dequa te a rm c i r cum f e r ence s . L e rbe rghe ( 1 988 ) s um s u p h i s r e cen t r e v iew o f t h e r e la t ion sh ip b e twe en c h i ldhood s t un t ing a nd m o r ta l i ty i ns t a t ing : " I f b o th s t un t ing a nd mo r ta l i ty a r ed i f f e r en to u t come s .o f as u cce s s ion o fs t re s se so nc h i ld r en ' sh ea l th , t h en t h e m ea su r e o f t h e h e igh t o f c h i ld r en i na c ommun i ty i sa n e ven m o re v a luab le t o o l f o r e va lua t ion . " ( 1 988 :259 ) . H e t h en g oe so n t os a y : " . . the m ea su remen t o f t h e p r eva lence o f s t un ted c h i ld r en t h en b ecome s a n o pe ra t iona l ly f a i r ly f e a s ib le , b e i ti n d i re c t , m ea su remen t o f p ove r ty r e la ted d i sea se f r equency . .T he p r eva len ce o f s t un ted c h i ld ren i n ac ommun i ty a ppea r s t ob e a g ood o ve ra l l i n d ica to r o f t h e h ea l th s t a tu s o f a c ommun i ty o fc h i ld ren . . ." ( 1 988 :259 ) . Ma r to re l l c ogen t ly s umma r ize s t h e p rob lem w i th t h es ma l lb u th ea l thy h ypo the s i sb ya s se r t ing t h e f o l low ing f o u r p o in t s . F i r s t , i n d iv idua l s , c h i ld r en a nd a du l t s , i nd eve lop ing c oun t r ie s a r e s ho r t d ue t oc h ron i ca l ly p oo r d i e t s a nd f r equen t i n f ec t ion d u r ing d eve lopmen t . T o c on s id e r s t un t ing t o b e a dap t ive i st o i n s inua te t h a t i t s c au se sa r ed e s i rab le . S e cond , g r ow th r e ta rda t ion , " r a the r t h an a n i n no cuou s r e spon se t o e nv i ronmen ta l s t imu l i , i s aw a rn ing s i gna l o f i n c r ea sed r i sk o f mo rb id i ty a nd m o r ta l i ty . " ( Ma r to re l l , 1 989 :15 ) . T h i rd ,t h ec ond i t ion sw h i ch c au se s t un t ing a l so a f f ec to t he rf u nc t ion s s u ch a s c ogn i t ion a nd w o rk c apa c i ty . F i na l ly , s t un ted g i r l s w ho g r ow u p s ho r t a r e a ti n c r ea sed r i sk o f d e l ive r ing g r ow th r e ta rded i n f an t s w i th ag r ea t e r c hance o fd y ing ( Ma r to re l l , 1 989 :15 ) . I n c on s ide r ing h ow S ek le r c ou ld h ave m ade s u ch an ea r s igh t ed i n t e rp r e ta t ion o f t h e a dap t ive c on sequen ce s o fs t un t ing , P e l to a nd P e I to ( 1 989 ) s u gge s t t h a t S e lde r f e l l v i c t im t o w ha t t h ey i d en t i f y a st h e " qu i t t ing e a r ly " p rob lem ( a l so s e e H a r r i s , 1 988 ) . T hey p ropo se t h a t m any s o c ia l s c ien t i s t s a r e g u i l ty o f f r equen t ly q u i t t ing e a r ly , b e fo r e t h ey h ave e va lua ted a l l e f f ec t s , a nd e s pec ia l ly t h el o ng te rm c au se sa nd c on sequence s o f " a dap ta t ion s . " S ek le r i s c l ea r ly g u i l ty o f q u i t t ing e a r ly . F u r the r , t h e c l a im o f a" n o -co s t a dap ta t ion " ( Bea ton , 1 989 ) s u gge s t s t o o m uch o f a na pp rova lo ft h es t a tu sq uo . Wha t a r e t h e i mp l ica t ion s o f t h e s t udy o f s t un t ing a nd t h e d i sp rov ing o f t h e " sma l l b u t h ea l thy " h ypo the s i s f o r s t ud ie s o fa dap ta t ion a nd h ea l th i np a s ts o c ie t ie s ? T he i mp l ica t ion st h a tc a n b ed r awn f r om d a ta o n t h ep r eva lence o fs t un t ing i nc on tempo ra ry p opu la t ion s s ugge s t st h a ts im i la r d a ta m igh t a l so b e ap owe r fu l i n d ica to ro fg r oup a d ju s tmen t i n p r eh i s to r i c s o c ie t ie s . We a nd o t he r s h ave n o ted s ome o f t h e d i f f icu l t ie s i n s t udy ing g r ow th p a t te rn s f r om s k e le ta l r ema in s ( Goodman e ta l . , 1 984 ; L a r sen , 1 9 87 ) . S ome o f t h e se p r ob lem s i n c lude f r agmen ta ry a nd s ma l l s amp le s i ze s a nd a n i n ab i l i ty t o p r ec i se ly a ge i n d iv idua l s . O nt h eo t he rh and ,l o ng b one l e ng th s c an b e a c cu ra te ly m ea su red a nd c ompa red t oa m ode rn s t anda rd ( Ma r e sh , 1 9 55 ) . I n f e rence s c an

b em ade f r om t h e s imp le p r eva lence o fs u badu l t s a nd a du l t w ho f a l l , f o r e xamp le , t wo s t anda rd d ev ia t ion so rg r ea te rb e low t h em ean .O nb a lance , t h e n umbe r o fc l ea r a nd p owe r fu l i n f e r en ce s t h a t c an b e g a ined f r om a n a na ly s i s o fg r ow th s t a tu s f a ro u twe igh s t h e t e chn i ca l d i f f icu l t ie s ( M ar t in e t a l . , 1 990 ) . E xam ined i nt h ec on tex to ft h e mode l ( F igu re 4 . 2 ) a nd t h e d a ta o n t h e c a u se a nd c on sequence s o f s t un t ing i n c on t empo ra ry p opu la t ion s , g r ow th d a ta c an b e u s ed t o m ake s t rong i n f e r en ce sa bou tg r oup l e ve la dap ta t ion . T he m o r e g ene ra l i mp l ica t ion t h a t c an b e d e r iv ed f r om t h e" sma l lb u th ea l thy "d eba t ei st h a t m any p a leopa tho log i s t s a l so q u i t t o o s o on . We q u i t b e fo r e s y s tem ica l ly a na lyz ing t h e i mp l ica t ion so fi n d iv idua ld i sea se p a t t e rn so n t h e g r oup a nd s o c i e ty . I n ac on tempo ra ry c on tex t , C havez a nd Ma r t inez ( 1 982 ) h ave s h own t h a t m ode ra t e ly m a lnou r i shed c h i ldren m ay b es i ck a s m uch a sh a l ft h et i me d u r ing t h e i rc h i ldhood . I ti s d i f f icu l t t oc omp rehend t h e b u rden t h i s l e ve l o f i l n e s s m u s t p l a ce o n am o the r i n ad eve lop ing c ommun i ty ( L ea the rman , 1 987 ) , a nd t h e r e i sn o r e a son t os u gge s t t h a t t h i s b u rden o fi l ne s s w a s a ny l e s s i nm o s t p a s t s o c ie t ie s . T oo o f ten w e i g no r e q ue s t ion s a bou t t h e b u rden o f i l ne s s o n t h e f u nc t iona l a nd a dap t ive c apa c i t ie s o f i n d iv idua la nd g r oup s . O r tne r ( 1 990 ) h a s v o iced o ne i n t e r e s t ing c ha l lenge t on o t q u i t t ing e a r ly . H i s c once rn i s t h a t w e t h ink c a r e fu l ly a bou t t h e m ean ing o f i n f ec t iou s l e s ion s o n b one . O r tne r s u gge s t s t h a t t h e se l e s ion ss hou ld n o tb ei n t e rp re ted a s as i gn o f s t r e s s ,b u tr a the ra ss i gn so fr a l ly ing f r om i n su l t s , i n d ica t ing a na dap t ive r e spon se t oi n f e c t ion . J u s t a s t h e r a i s ing o f t h e q ue s t ion w a s i mpo r tan t f o r n u t r i t ion i s t sw ho r a l l ied t oc ons ide rt h ev a l id i ty o f t h e " sma l l b u t h ea l thy " h ypo the s i s , t h e r a i s ing o f t h eq ue s t ion c once rn ing t h e a dap t ive s i gn i f i cance o fs k e le ta l i n d i ca t ion s o fd i sea se i sa n i mpo r tan t c a l l t o p a le ,opa tho log i s t t o t h ink t h rough t h e i r d a ta . Is u gge s tf u r the rt h a tO r tne r ' sc ha l lenge h a s t og oe ven f u r the rt on o tj u s tt h ink ing t h rough t h e i n d iv idua lr e spon se t ot h ei n su l t ,b u tt h e s y s tem ic e f f ec t t h a t t h e s t rugg le t oa dap t t oa n i n f ec t iou s d i sea se m igh t h ave o n af am i ly a nd l a rge r s o c ia l g r oup s . A n umbe r o f r e sea r ch i n i t ia t ives c an s t r eng then t o ag r ea te r e x t en t i n f e r en ce s o n t h e f u nc t iona l a nd a dap t ive c on sequence s o f o u r i n d ica t ion s o fh ea l th . I ns ome c a se s i tm igh t b e i mpo r tan t t o k now t h e c onsequence s o f a n i n d ica t ion o f s t r e s s i n ac on tempo r a ry c on t ex t . A n e xamp le o ft h i s c once rn s e name l h ypop la s t i c d e f e c t s . I no r de rt om ake c l ea re ri n f e r enc es a bou t t h e m ean ing o fe name l h ypop la s ia s , w e n eed t o b e t t e rk now t h e s e n s i t iv i ty a nd s p ec if i c i ty o ft h i s i n d ica to r t ob ou t s o f m a lnu t r i t ion a nd i n f ec t ion ( Goodman a nd R o se , 1 990 ) . I no t he rc a se s t h e u nde r ly ing c au se a sw e l l a s t h e f u nc t iona l c on sequence s o f as t r e s so r a r e k nown . F o re xamp le ,i ti sc l ea rt h a tm o s tc a se so f p o ro t ic h ype ro s to s i s a r e s e conda ry t o i r on

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d ef ic iency , a nd t h e e f fe c t s o f i r on d ef i c ien cy o n f u nc t ion a r e i n c rea s ing ly k nown f r om c on t empo ra ry s t ud ie s ( Chwang e t a l . , 1 988 ; D a l lman , 1 987 ; P o l l i t t , 1 987 ) . I n t h i s c a se , r e sea r ch i s n eeded o n t h e p re c i se r e la t ion sh ip b e tween t h e s k e le ta l i n d ica to r s o fi r on d e f i c iency a nd b i ochem i ca l i n d i ca to r s . H ow c h ron i c a nd s e ve r e a d ef i c iency i s i n d ica t ed b y i r on d e f ic iency? A l though s im i la r k now ledge " g ap s " a r e f o und f o r mo s t p a leopa tho log i ca l m ea su r e s , I r ema in o p t im i s t ic a bou t t h e a b i l i ty o f w e l l d i re c t ed r e sea r ch e f fo r t st of i l lt h e se g ap s .

C onc lus ion sa nd R ecommenda t ion sf o r F u ture R e search T he re a r e t h r ee f u ndamen ta lp o in t s t h a t Ih ope t o h ave u nde r sco red i n t h i s d i s cu s s ion o f o u r a b i l i ty t ou nde r s tand h ea l th a nd i t sr e la t ion sh ip t o a dap ta t ion i np a s ts o c ie t ie s . T he m o s tg ene ra la nd t h eo re t ica l p o in t c once rn s t h e s yne rge t ic a nd i n t e ra c t ive r e la t ion sh ip b e tween h ea l th a nd a dap ta t ion . H ea l th i s af u nc t ion o ft h eo r gan i sm ' s a b i l i ty t oa d ju s t t oe nv i ronmen ta l c on s t r a in t s a nd s t re s so r s . I ti st h e r e su l to ft h e a dap t ive p ro ce s s . P oo r h ea l th c on s t ra in s a dap ta t ion . I t i s a c ond i t ion t h a t n e ed s t ob e a lev ia ted o r a d ju s t ed t o . I n t h i s s e n se , h ea l th , c l a s s i ca l ly d e f ined i n t e rms o fm o rb id i ty a nd m o r ta l i ty , i s am ea su r e o f t h es u cce s so fa dap ta t ion . U nde r f a vou rab le c ond i t ion s , a dap ta t ion i s m an if e s t i ng ood h ea l th , a nd h ea l th f u r the r s o ne ' s a b i l i ty t o a d ju s t t o s t re s s f u l c ond i t ion s . S uch c ond i t ions c an i n c rea se o ne ' s a dap t ive f l ex ib i l i ty a nd a b i l i ty t o e ndu r e o t he r s t r e s so r s a nd c ons t ra in t s . C onve r se ly , t h e i n ab i l i ty t oa dap t i s a ppa ren t i np oo r h ea l th , w h ich i nt u rn d e c rea se s o ne ' s a b i l i ty t oa dap t t of u r the r i n su l t s . S t r e s s f u l e nv i ronmen ta l c ond i t ion s p romo t e h ea l th p rob lem s , w h ich c on s t ra in a dap ta t ion b y , a mong o t he r t h ing s , l owe r ing r e s i s tan ce t o d i sea se , l owe r ing w o rk a nd p roduc t ive c apa c i ty , l owe r ing i n t e r e s t i n a nd a b i l i ty t o e ngage i n s o c ia l a nd d i s c re t iona ry a c t iv i t ie s , i n te r f e r ing w i th l e a rn ing a nd c ogn i t ive f u nc t ion ing ,d ec r ea s ing f e r t i l i ty a nd f e cund i ty , a nd i n te r f e r ing w i th n u t r ien ti n take a nd u t i l iza t ion ( A l len , 1 984 ; B uz ina e ta l . , 1 989 ) . T he p ro ce s sb y w h ich h ea l th i sr e la t ed t oe nv i ronmen t a nd a dap ta t ion i s n o th ing l e s s t h an i n t r in s i c , i n t e ra c t ive ,a nd c yc l ica l . T he s e cond f u ndamen ta l p o in ti st h a th ea l th i s am ea su rab le e n t i ty i ns t ud ie s o fc on t empo ra ry a s w e l la sp a s ts o c ie t ie s . R ead i ly a va i lab le m ea su r e s o fh ea l th s t a tu s , s u ch a sr a t e s o f i n f an t m o r ta l i ty a nd s t un t ing , a r e r ou t ine ly u s ed t o g auge a nd c ompa r e t h e g ene r a l l e ve l o f h ea l th o f c on t empo ra ry g r oup s ( Goodman e t a l . , 1 988 ) . Wh i le h ea l th s t a tu s , a s w e l l a s s t r e s s a nd n u t r i t iona l s t a tu s , i s ab ehav ior a l p henomena t h a t c an n o t b e d i r ec t ly m ea su r ed i n p a s t s o c ie t ie s , t h e re a r e n ume rou s c ommon ly e mp loy ed a nd i n c rea s ing ly r e f ined m e thod s f o r r e con s t ruc t ing h ea l th , s t re s s , a nd n u t r i t iona l s t a tu se s f r om b one s

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a nd t e e th ( s e e f o r e xamp le s t h e f o l low ing r e v iew s : B u ik s t ra a nd C ook , 1 980 ; G oodman e t a l . , 1 984 ;G i lbe r ta nd M ie lke ( e d s . ) , 1 985 ;L a r sen , 1 987 ) . S ome o f t h e se m ea su remen t s , s u ch a s t h o se i n vo lv ing mo r ta l i ty a nd g row th r a t e s , a r e p a r t icu la r ly r obu s t i n t h e i r a b i l i ty t o g ene ra te a dap t ive i n f e r ence s b ecau se t h e re a r e e qu iva len t a nd c ommon ly u s ed c on tempo ra ry m ea su re s ( Goodman e ta l . , 1 988 ) . Wh i le t a phonom ic p ro ce s se s r e nde r i t i mpo s s ib le t or e con s t ruc t f u l ly a l d imen s ion s o f h ea l th e xpe r ien ced b y a n i n d iv idua l o r ag roup e i the r i n t h e p a s t o r c u r ren t ly - I s ubm i t , o p t im i s t ica l ly b u t n o t w i thou t e mp i r ica l a nd s c ien t i f ic j u s t if ica t ion , t h a t m any t e chn ique s a r e n ow a va i lab le t h a t c an p rov ide a n a ccu ra t e a pp ra i sa l o f an umbe r o fd imen s ion s o f h ea l th i n p a s t p opu la t ion s . T he i n fo rma t ion d e r ived f r om t h e se t e chn ique s , c oup led w i th a n u nde r s tand ing o ft h e a dap t ive a nd f u nc t iona l s i gn if icance o ft h e m ea su r e s o fh ea l th , c an p rov id e k ey i n s igh t s i n to t h ed ynam ic so fa dap ta t ion i np a s ts o c ie t ie s . T h i rd ly , s t ud ie s o fh ea l th i np a s t s o c ie t ie s c an p rov ide u n ique i n s igh t s i n to t h e d ynam ic s o fp a s t s o c ie t ie s a nd t h e e vo lu t ion o f t h e h uman c ond i t ion . U n fo r tuna te ly ,m any p a leopa tho log i s t s h ave q u i t t o o s oon , b efo r e t h e s i gn if icance o f h ea l th h a sb een f u l ly e xp lo r ed . I nc onc lud ing Iw i sh t om ake r e f e r ence t ot h e t h eo r e t ica l u nde rp inn ing o f t h e m ode l a nd p e r spe c t ive o u t l ined a bove i n r e la t ionsh ip t o r e cen tt h eo r e t ica l c ha l lenge s i na r chaeo logy . I ti s s omewha t p a radox ica l t h a t t h e r e sea r ch p a rad igm w h ich i s h i gh ly p ro ce s sua l i s t i n n a tu re i s p ropo sed a t at i me w hen a r chaeo logy s e ems t ob e m ov ing a way f r om ap roce s sua l is t p a r ad igm ( f o r e xamp le s e eH odde r , 1 986 ) . T h i sp a radox m ay b e e xp la inab le b y t h e f a c t t h a t o ne o f t h e p ro ce s sua l i s t p a rad igm ' s g r ea te s t f a i l ing s , l a ck o f c once rn f o r i n d iv idua l a nd c u l tu ra l ly s pe c i f ic d ynam ic s , i s f a r l e s s a pp l icab le t o b i o log ica l p henomena . T he b i o log i ca l i mpa c t a nd c on sequen ce s o fh ea l th a r e m o re n ea r ly u n ive r sa l t h an s o c ia l p ro ce s se s . T he re f o re , c on tempo ra ry a na log ie s a bou t t h e c ause a nd e f f ec t o fh ea l th a r e q u i t e c on s i s ten t a nd c an b e u s ed t o m ak e a s sump t ion s a bou t c au se a nd e f f ec t i n t h e p a s t . F u r the r , a n e xam ina t ion o fh ea l th c an a nd s hou ld b es e en n o ta si s o la t ed l o ca l leve lp henomena ,b u t a s ar e su l t o f t h e o penne s s o f l o ca l s y s tem s w i th o t he r s y s t em s . I n f a c t , t h e mode rn e t hnog raph ic d a ta s ugge s t t h a tp oo r h ea l th i si n va r iab ly t r a cked b ack t op ove r ty , w h ich i si t se lf d ue t or e g iona l a nd i n te r r eg iona lp ro ce sse s . T he f a u l to ft h e n ew a r chaeo logy o fo ve re mpha s i s o n l o ca l e co log ica l p ro ce s se s n e ed n o t b e s ha red b y ap ro ce s sua l i s t v i ew o f h ea l th a nd a dap ta t ion . T hu s , i ti sn o t p a radox i ca l t h a t w hen s ome s o c ia l a r cha eo log i s t s a r e a bou t t o d i sca rd p ro ce s se s , m idd le r a nge t h eo ry , a nd e t hnog raph i c a na logy , b i oan th ropo log i s t sm ay d i scove rt h e u s e f u lne s so f t h e se m e thodo log ica l a nd t h eo r e t ica l u nde rp inn ing s . T he s t udy o f h ea l th i n p a s t

s o c ie t ie s n eed s t o b e p l a ced i na n e vo lu t iona ry a nd e co log ica l c on tex t , l i nk ing i tt o s t ud i e s o f c on t empo ra ry g r oup s . I nt h i s w ay w e w i l l b e t t e r u nde r s tand i d ea s a bou t p rog re s s , t h ed ynam i c s o f p a s t s o c ie t ie s , a nd t h e e vo lu t ion o f t h e h uman c ond i t ion .

H a r r i s , M . ( 1 988 ) . C u l tu ra l m a te ra l i sm : a l a rum s a nd e xcu r s ion s . I n Wayma rk s : T he N o tre Dame I naugura lL ec ture s i nA n th ropo logy . e d . K . Moo re . p p . 1 07 -26 . N o t re D ame : U n ive r s i ty o fN o t re D ame P r e s s . H odde r , I .( 1 986 ) . R ead ing t h e P as t . N ew Y o rk : C amb r idge U n ive r s i ty P re s s . H u s s -A shmo r e , R . , G oodman , A . & A rme lago s , G . ( 1 982 ) . N u t r i t iona l i n f e rence f r om p a leopa tho logy . I nA dvance s i nA rchaeo log ica l Me thod a nd T heo ry V o l . 5 . e d . M . B . S ch if f e r . p p . 4 36 -74 . O r lando : A cadem ic P re s s . L a r sen , C . ( 1 987 ) . B ioa rchaeo log ica l i n te rp re ta t ion s o f s ub s i s tence e conomy a nd b ehav iou r f r om h uman s k e le ta l r ema in s . I n A d vances i n A rchaeo log ica l Me thod a nd T heo ry V o l . 1 0 . e d M . B . S ch i f f e r . p p . 3 39 -445 . O r lando : A cadem ic P re s s . L ea the rman , T . ( 1 987 ) . I lne ss , Work a nd S oc ia l R e la t ions i n t h e s ou the rn P eruv ian H igh land s . U npub l i shed D oc to ra l D i s se r ta t ion , U n ive r s i ty o f Ma s sachu se t t s , Amhe r s t , D epa r tmen t o f A n th ropo logy . L e rbe rghe , W . ( 1988 ) . L inea r g row th r e ta rda t ion a nd m o r ta l i ty . I n L inea r G row th R e tarda t ion i n L e ss D eve loped C oun t r ie s . e d . J .C . Wa te r low . p p . 2 45 6 4 . N ew Y o rk :R aven P re s s . L iv ing s tone , F . ( 1 958 ) . A n th ropo log ica l i mp l ica t ion s o f s i ck le c e l l g ene d i s t r ibu t ion i n We s t A f r ica . Am . A n thropo l . 6 0 :533 -62 . Ma re sh , M . ( 1 955 ) . L inea r g r ow th o f l o ng b ones o f e x t r em i t ie s f r om i n f ancy t h rough a do le scence . A n t . J .D i s .C h i ld . 8 9 :725 -42 . Ma r t in , D . , G oodman , A . , A rme lago s , G . & Magenn i s , A . ( 1 990 ) . B lack Me sa A na sa z i H ea l th : R econ s truc t ing L tf e f rom P a t te rns o f D i sease a nd D ea th . C a rbonda le : S ou ther n I l ino i s U n ive r s i ty P re s s( I np r e s s ) . Ma r to re l l , R . ( 1 980 ) . I n te r re la t ionsh ip b e tween d i e t , i n f ec t iou s d i sea se , a nd n u t r i t iona l s t a tu s . I n S oc ia l a nd B io log ica l P red ic to r s o f N u t r i t iona l S ta tus , P hys ica l G row th a nd N euro log ica l D eve lopmen t . e d s . L . G reene a nd F . J ohn s ton . p p . 8 1-106 . O r lando :A cadem ic P r e s s . Ma r to re l l , R . ( 1 989 ) . B ody s i ze , a dap ta t ion a nd f u nc t ion . H um .O rgan .4 8 :15 -20 . Ma ta ,1 . ,U r ru t ia ,J .& L ech t ig , A .( 1971) . I n f ec t ion a nd n u t r i t ion o f c h i ld r en o f al ow s oc ioeconom ic r u ra l c ommun i ty . Am .J .C l in .N u tr .2 4 :249 -59 . May , J .( 1 960 ) . T he e co logy o fh uman d i sea se . A nn . N .Y .A cad .S c i .8 4 :789 -94 . McKeown , T .( 1 988 ) . T he O r ig ins o fH uman D i sease . N ew Y o rk : B lackwe l l . O r tne r , D . ( 1 990 ) . T heo r e t ica l a nd m e thodo log ica l i s sue s i np a leopa tho logy . I nH uman P a l eopa tho logy : C ur ren t S yn the se s a nd F u ture O p t ion s . e d s . D . O r tne r a nd A . A u f te rhe ide . Wa sh ing ton , D .C . : Sm i th son ian I n s t i tu t ion P re ss ( I np re ss ) . P e l to , G . a nd P e l to , P .( 1989 ) . Sma l l b u t H ea l thy? A n A n th ropo log ica lP e r sp ec t ive . H um .O rg .4 8 :11 -15 . P o l l i t t , E .( 1 987 ) . E f f ec t s o fi r on d e f ic iency o n m en ta l d eve lopmen t : m e thodo log ica l c on s ide ra t ions a nd s ub s tan t ive f i nd ing s . I n N u t r i t iona l A n th ropo logy . e d . F . J ohn s ton . p p . 2 25 -54 . N ew Y o rk : A lan R . L i s s , I n c . S ek le r , D . ( 1 980 ) . Ma lnu t r i t ion : a n i n t e l lec tua l o dy s sey . W e s t .J .A gr ic .E con .5 :219 -27 . S ek le r , D . ( 1 982 ) . Sma l l b u t h ea l thy? S ome b a s ic p rob lem s i n t h e c oncep t o f m a lnu t r i t ion . I n N ewe r C oncep ts i n N u t r i t ion a nd T he i r I mp l ica t ion s f o r

R e f erence s A lam , N . , Wo j tyn iak , B . & R ahaman , M . ( 1 989 ) . A n th ropome t r ic i n d ica to r s a nd r i sk o f d ea th . Am . J . C l in .N u t r .4 9 :884 -88 . A l len , L . ( 1984 ) . F unc t iona l i n d ica to r s o f n u t r i t iona l s t a tu s o f t h e w ho le i nd iv idua l o r t h e c ommun i ty . C l in .N u t r .3 5 :169 -75 . B ea ton , G . ( 1989 ) . Sma l lb u tH ea l thy? A re w e a s k ing t h er i gh tq ue s t ion? H um .O rg .4 8 :30 -39 . B u ik s t ra , J .a nd C ook , D . ( 1 980 ) . P a leopa tho logy : a n Ame r ican a c coun t . A nn .R e v .A n thropo l .9 : 433 -70 . B uz ina , R . , B a te s , C . , v an d e r B eek , J . , B rubache r , G . , C hand ra , R . , H a l lb e rg , L . , H e sek e r , J . , Me r tz , W . , P i e t rz ik , L . , P o l l i t t , E . , P rad i l la , A . , S ubo t icanec , K . , S and s tead , H . , S cha lach , W . , S pu r r , G . & We s tenhof e r , J . ( 1 989 ) . Wo rk shop i n f unc t iona l s i gn if i cance o f m i ldt o -mode ra te m a lnu t r i t ion . Am . J .C l in .N u tr .5 0 :172 -76 . C havez , A . a nd Ma r t inez , C . ( 1 982 ) . G row ing U p i na D eve lop ing C oun t ry . Mex ico : I n s t i tu to N ac iona l d e l aN u t r ic ion . C hen , L . , C howdhu r ry , A . , & H of fman , S . ( 1 980 ) . A n th ropome t r ic a s se s smen t o f e ne rgy -p ro t e in m a lnu t r i t ion a nd s ub sequen t r i sk o fm o r ta l i ty a mong p r e schoo la ged c h i ld r en . Am .J .C l in .N u t r . ,3 3 :1836 4 5 . C hwang , L . , S oema r t r i , A . &P o l l i t t , E . ( 1 988 ) . I r on s upp lemen ta t ion a nd p hy s ica l g row th o f r u ra l I n done s ian c h i ld ren . Am .J .C l in .N u tr . 4 7 :496 -50 1 . D a l lman , P . ( 1987 ) . I r on d ef ic iency a nd t h e i mmune r e spon se . Am .J .C l in .N u tr . 4 6 :329 -34 . D ie tz , W . ( 1 983 ) . O ne f o r a l? t h ough t s o n r e f e r ence s t anda rd s f o r g r ow th i ns ho r tp opu la t ion s . N u tr . R e s . 3 : 129 -31 . G i lb e r t ,R . a nd M ie lke , J .( 1 985 ) . e d s . T he A na l ys i so f P reh i s to r ic D ie ts . O r lando : A cadem ic P re s s . G oodman , A . a nd A rme lago s , G . ( 1 989 ) . I n f an t a nd c h i ldhood m o rb id i ty a nd m o r ta l i ty r i sk s i n a r chaeo log ica l p opu la t ion s . Wo r ld A rchaeo l . 2 12 :225 -43 . G oodman , A . , Ma r t in , D . , A rme lago s , G . &C la rk , G . ( 1 984 ) . I nd ica t ion s o f s t r e s s f r om b one s a nd t e e th . I n P a l eopa tho logy a tt h eO r ig ins o fA g r icu l tu re . e d . M .N . C ohen a nd G .J . A rme lago s . p p . 1 3 -49 . O r lando : A cad em ic P r e s s . G oodman , A . a nd R o se , J .( 1 990 ) . D en ta l e name l h ypop la s ia s a s i n d ica to r s o f n u t r i t iona l s t a tu s . I n A dvance si nD en ta lA n thropo logy . e d s . M . K e l ly a nd C .L a r sen . AR L i s s : N ew Y o rk ( I np re s s ) . G oodman , A . , T homa s , R . , S wed lund , A . & A rme lago s , G . ( 1988 ) . B iocu l tu ra l p e r sp ec t ive s o n s t re s s i n p r eh i s to r ic , h i s to r ica l , a nd c on tempo ra ry p opu la t ion r e sea rch . Y ea rb . P hy s . A n th ropo l . 3 1 :169 -202 . G ra i tce r ,P .a nd G en t ry ,E .( 1981) . Mea su r ing c h i ld r en : o ne r e f e rence f o ra l . L ance t2 : 297 -99 . H ab ich t , J . -P . , Ma r to re l l , R . , Y a rb rough , C . , Ma l ina , R . &K le in , R . ( 1 974 ) . H e igh ta nd w e igh t s t anda rd s f o r p r e schoo l c h i ld ren . H ow r e levan t a r e e t hn ic d i f f e rence s i ng r ow th p o ten t ia l? L ance t1 : 611 -15 .

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P o l icy . e d . P .V . S u l cha tme . p p . 1 39 -48 . P une , I nd ia : Maha ra sh t ra A ssoc ia t ion f o r t h e C u l t iva t ion o f S c ience . Smedman , L . , S t e rky , G . , Me l land e r , L . & Wal , S . ( 1 987 ) . A n th ropome t ry a nd s ub sequen t m o r tal i ty i n g r oup s o f c h i ld ren a ged 6 59 m on th s i n G u inea B i s sau . Am .J .C l in .N u t r .4 6 :369 -73 .

F igure 4 . 1

T homa s , R .B . ,W in te rha lde r ,B . & McRae , S .D . ( 1 979 ) . A n a n th ropo log ica l a pp roach t oh uman e co logy a nd a dap t ive d ynam ic s . Y ea rb .P hys .A n thropo l .2 2 :1 -46 . T ompk in s , A . ( 1 988 ) . T he r i sk o f mo rb id i ty i n t h e s t un t ed c h i ld . I nL inea r G row th R e tarda t ion i nL es s D e ve loped C oun t r ie s . e d . J . C . Wa te r low . p p . 1 85 9 9 . N ew Y o rk :R aven P re s s .

Ap ea san tf am i ly f r om T ezon t eopan , Mex ico . T he t wo o l de rg i r l s a r ee n ro l led i nt h e s t udy o fC havez a nd Ma r t inez ( 1 982 ) . T he o ne w i th t h e w h i te t sh i r t w i th t h e r o se ( Ro sa ) i sn i ne t een a nd w a s n o t n u t r i t iona l ly s u pp lemen ted . T he o t he r s i s te r i sf i f teen y ea r s o l d a nd h a s b een g i ven n u t r i t iona l s upp lemen t s s i nce b i r th . T he f i ve i n ch h e igh td i f f e r ence b e twe en t h e se s ame s e x s i b l ing i st y p ica l o f t h e d i f f e rence s b e tween t h e s upp lemen ted a nd n on supp lemen ted c h i ld r en . I n a dd i t ion t ot h e h e igh t d i f f e rence s , C havez a nd Ma r t inez ( 1 982 ) h ave f o und p rof ound d i f f e rence s i n a c t iv i ty l e ve l s , s oc iab i l i ty ,a nd r e s i s tance t od i sea se .

3 7

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D IRECT GENET IC AND IM MUNOLOG ICAL I NFOR MAT ION I N THE RECONSTRUCT ION OF HEALTH AND B IOCULTURAL COND IT IONS OF PAST POPULAT IONS : A NE W PROSPECT FOR ARCHAEOLOGY .

C r i s t ina C a t taneo t h e e nv i ronmen t s u r round ing h im . I t i s a l so i mpo r tan tt oa cqu i r e ap rop e ru nde r s tand ing o ft h e c au sa t ive f a c to r s b eh ind g ood o r b ad h ea l th c ond i t ion s . F u r the r , i ti sc r uc ia l t o r e a l i se t h a t t h e re a r e t wo e qua l ly r e levan t m a in c on t r ibu t ing f a c to r s t od i sea se : o ne i st h e e nv i ronmen t , t h e o t he r ,n o tl e s si mpo r tan t ,i sg ene t ic p r ed i spo s i t ion t od i sea se ( F igu re 5 . 1 ) .

H ea l th a nd A rchaeo log ica l R econs truc t ion I nr e cen t y ea r s , t h e i n c rea s ing i n te re s t i nt h e s t udy o f o rgan i c r ema in s ( b o tan i ca l , f a una l , h uman ) r e t r ieved f r om a r chaeo log ica l c on tex t s h a s b rough t f o r th a s e r iou s a t t emp t t o i n t eg ra te i n fo rma t ion o b ta ined f r om b i o log ica l i n ve s t iga t ion s w i th t h a t o f a mo r e t y p i ca l ly a r chaeo log ica l n a tu re s u ch a s , f o r e xamp le , m a te r ia l c u l tu r e . H oweve r , r e ga rd le s s o f t h e se n ew a pp roache s , t h e b i omed ica l f i e ld h a s n o t y e t b een a b le t o f i nd i t s d e se rved n i che w i th in t h e d i s c ip l ine o fa r chaeo logy . P r ev iou s s t ud ie s i np a la eopa tho logy h ave , i n f a c t , c on f ron t ed t h e c ha l lenge s o fu n rave l l ing t h e h i s to ry a nd e vo lu t ion o f s pe c i f ic d i sea se s , o f d i agno s ing p a tho log ica l c ond i t ion s i ni n d iv idua l s k e le tons , a nd o f u s ing t h e m o s t mod e rn b i omed ica l l a bo ra to ry t e chn ique s t o e x t ra c t b i omo lecu le s s uch a s h aemog lob in , a l bum in , c o l lagen a nd DNA f r om h uman s k e le ta l m a t e r ia l . A l l t h e se m ay b e v e ry i n te re s t ing a ch ievemen t s p e r s e b u t , w hen i s o la ted f r om a b i o cu l tu ra l c on t ex t , t h ey a r e u s e le s s i nt h e r e con s t ru c t ion o f l i f e s ty le a nd d ynam ic so fp a s tp opu la t ion s . T he s t a te o f h ea l th i n p a r t icu la r s hou ld b e c on s ide red c r uc ia l i n f o rma t ion b y t h e a r chaeo log i s t , e spe c ia l ly w hen a s se s sed a t a p opu la t ion l e ve l , f o r i ti si n t e r tw ined t i gh t ly w i th p henomena s u ch a sp opu la t ion d ynam i c s ,l i f e s ty le a nd c u l tu r ec hange . F rom t h eh ea l th c ond i t ion so f a p opu la t ion o r o f s ubg roup s a mong t h a t p opu la t ion , o ne c an b eg in t o m ake i n f e r en ce s c oncer n ing s ub s i s ten ce , n u t r i t ion , c l a s s d i f f e r ence s , a nd s o f o r th . F o r e xamp le , w hen a mong a s ke le ta l p opu la t ion c e r ta in s ubg roup s s e em t oh ave b een a f f ec t ed ( t o ag r ea t e r d eg r ee t h an o t he r g roup s ) b y " s k e le ta l ly d i agno sab le " s t r e s s f a c to r s , o r s e em t o h ave s u f f e r ed f r om n u t r i t iona l d e f ic ienc ie s , t h ough m a te r ia l c u l tu re m ay n o tg i ve u sa ny c l ue sc once rn ing t h i sm a t t e r , w e c an , o n t h e b a s i s o f s k e le ta l e v idence o n ly , b eg in t o f o rmu la te t h e h ypo the s i s t h a t t h e se i n d iv idua l s b e longed t o al e s s p r iv i leged s o c ia l c l a s s . I n o t he r w o rd s , ac l o se ly k n i t i n t e ra c t ion b e tween t h e s t udy o f c u l tu r e a nd t h a t o f h uman b i o logy i se s sen t ia l . T o r e con s t ruc t t h e h ea l th s t a tu s o f a p opu la t ion i ti sf i r s t n ece s sa ry t ou nde r s tand t h e i n t e ra c t ion b e tween m an a s ab i o log ica le n t i ty a nd

H ea l th : G ene t ic so rE nv ironmen t? U nde r t h e h ead ing " e nv i ronmen t " i t i s r e a sonab le t o i n c lude a n a r ray o f p henomena w h ich m ay i nf luence t h e i n d iv idua l f r om t h e " o u t s id e " . T h i s a r ray m ay r a nge f r om m i c ro o r gan i sm s , af r equen tc au se o fd i sea se , t oc l ima t ic c ond i t ion s , e conomy a nd e ven c u l tu ra l h ab i t s , w h ich m ay a f f ec t t h e l i f e s ty le o f a n e n t i r e p opu la t ion ( ac e r ta in d eg re e o f c i r cu la r i ty i s , o f c ou r se , i n ev i tab le w hen t h e r e c ip ro ca l e f f e c t s o f h ea l th /b io logy a nd c u l tu r e a r e c on s ide red ) . O n t h e o t he r h and t h e t e rm " g ene t i c p r ed i spo s i t ion " r e f e r s t ot h e i n na te p r ed i spo s i t ion o f i n d iv idua ls ( o r p opu la t ion s ) t od eve lop a p a r t icu la r d i sea se . K now ing t h e n a tu r e o f t h e se p r ed i spo s i t ion s i s e x t r eme ly a dvan tageou s , s i nce a l d i sea se s a r i se a s a r e su l t e i the r o f g ene t ic s u scep t ib i l i ty , e nv i ronmen ta l f a c to r s o r , m any t i me s , o f t h e se t wo p henomena c omb ined . I t i s p r ec i se ly t h e g ene t ic c omponen t o f d i sea se t h a t Iw i sh t oe l uc ida te i nt h i sa r t i c le . T he a im o ft h i s p ape r i st oe xp la in t h e r e levance , n o t o n ly i nt h e m ed i ca l b u ta l so i nt h ea r chaeo log ica l c on t ex t , o f t h e g ene t i c c on t r ibu t ion t o s ome d i sea se s ; t os how h ow t h i s i n fo rma t ion m ay h e lp u st or e con s t ruc tm o r ea c cu r a te ly t h eh ea l th s t a tu s a nd l i f e s ty le o f ap opu la t ion , a nd t os ugge s t h ow t h i s g ene t i c d a ta m ay b e o b ta ined f r om s p e c i f ic b i o log ica l m o le cu le s r e t r ieved d i rec t ly f r om s k e le ta lm a te r ia l( s ee b e low ) . S of a r , w e h ave s e en t h a t t om ake a n a c cu ra t e s t a temen t c on ce rn ing t h e l i f e s ty le a nd d ynam i c s o f ap opu la t ion o ne m u s t u nde r s tand i t s h ea l th s t a tu s , a nd t o d o t h i s , o ne m u s t r e a l i se h ow e x t en s iv e ly e nv i ronmen t a nd g ene t i c s c on t r ibu t e t od i sea se . T he r e fo r e ,t or e con s t ruc th ea l th s t a tu s , n o t o n ly s hou ld w e l o ok f o r t h e e nv i ronmen ta l f a c to r s w h ich m ay a f f ec t i t ,b u t w e m u s t a l so t r y t od e t e rm ine w ha ta nd h ow m uch w a sc on t r ibu ted b y g ene s a l one . T h i s k i nd o f i n fo rma t ion m ay h e lp a vo id i n comp le te o r e r roneou s r e con s t ruc t ion s o f h ea l th b a sed o n ly o n s k e le ta l

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l e s ion s a nd d emog raph ic p a t te rn s . I n o t he r w o rd s , o ne s hou ld t r y t o c omp rehend w he the r i n d iv idua l s w e r e d e s t ined , b e cau se o f s ome g ene t i c c ha r ac te r i s t i c o r a noma ly , t o d eve lop s p ec i f ic c ond i t ion s ( o r t ob e m o re s u s cep t ib le t o a cqu i r ing d i sea se s ) , o r i fe nv i ronmen ta l f a c to r s w e re t h e a gen t s c h ief ly r e spon s ib le f o r t h e o n se t o f ap a r t icu la r d i sea se . We s hou ld a vo id p i t f al s s u ch a s a t t r ibu t ing t h e c au se s o f i l ne s s o r b ad h ea l th t oe nv i ronmen ta l f a c to r s w hen t h e g ene t ic m ak eup i sp r ima r i ly r e spon s ib le . A n e xamp le o f t h i s k i nd o fe r ro r i ss e en w hen c on s ide r ing c a se s o f a naem ia d i agno sed a c co rd ing t o s k e le ta l l e s ion s . A t t i me s , t h e p a la epa tho log i s t i s c onf ron ted w i th s k e le ta l e v idence o f a naem ia i n t h e f o rm o f p o ro t ic h yp e ro s to s i s ( t h i ck en ing a nd p o ro s i ty o f t h e c r an ium a nd o f t h e r o o f o f t h e o r b i t s ) . O n o cca s ion , t h e se l e s ion s c ou ld h ave h ad ag ene t ic b a s i s ( a s i nt h e c a se o ft h a la s sem ia o r Med i te r ranean a na em ia ) . O the r t i me s , t h ey c ou ld h ave b een a c on sequence o f a dve r se e nv i ronmen ta l f a c to r s , s u ch a s i r on d e f i c iency . Many p a laeopa tho log i s t s ( e .g . S t e inbo ck , 1 976 ; O r tne r a nd P u t s cha r , 1 985 ) t r y t o d i f f e r en t ia te b e twe en t h e s k e le ta l m an i f e s ta t ion s o f i r on d e f ic iency a naem ia a nd t h o se o f g ene t i c a naem ia s . B u t t h i s i s ad i f f i cu l t t a sk , s imp ly b ecau se t h e c hange s i n t h e b one t i s sue a r e t o o n on spec if i c ( Ha r t . , 1 980 ) . I n p r a c t ice , t h e se l e s ion s o f t en h ave b een h a s t i ly a t t r ibu t ed t o d i e t a ry i r on d e f ic iency , a nd t h e h ypo the s i s t h a t t h e c ond i t ion m ay h ave b een g ene t ic i ss e t a s ide . T h i s t y pe o f e r ro r c ou ld b e e xcu sed i f t h e i mp l ica t ion s o f s u ch a d i agno s i s w e r e n o t s o i mpo r tan tt ot h ea r chaeo log i s t . I nf a c t ,a t t r ibu t ing t h e a naem ia t od i e ta ry d e f ic iency w i l l l e ad t ot h e i d ea t h a tt h ep opu la t ion ,o rp a r t s o fi t ,l i ved o n a n i n ad equa te d i e t , w he r ea s t h e d i sea se , b e ing g ene t i c ,m ay h ave b een i n ev i tab le ,e ven u nde rt h e b e s t o fn u t r i t iona l c ond i t ion s . A nge l ( 1 967 ) a l so h a s r e commend ed b e ing v e ry c a r e fu l o f t h i s p i t f a l l . I n s k e le ton s f r om t h e e a r ly N eo l i th ic ( c . 6 500 B .C . ) a nd B ronze A ge G r ee ce , o ne c an n o t i ce t h e se l e s ion s i nt h e b one s o fb o th c h i ld r en a nd a du l t s . A nge l h a sr e la ted t h i s t ot h ep r e sence o f e ndem i c m a la r ia a nd t h e r e fo r e , i n d i rec t ly , t o t h ep r e sence o ft h a la s sem ia . T ha la s sem i cs i nf a c t s e em t ob e p ro t ec ted , t o ac e r ta in d eg r ee , a ga in s t m a la r ia . A s e vo lu t iona ry t h eo ry w ou ld h ave i t , i n d iv idua l s c a r ry ing a " d eg r ee " o f t h a la s sem ia c ompa t ib le w i th l i f e w ou ld h ave b e en b e t t e r a dap ted a nd t h e re fo re s e lec ted f o r i n s u ch a n e nv i ronmen t . I n t h i s c on t ex t i ti sn o t p o s s ib le , a c co rd ing t o A nge l , t o m ake a ny d i e ta ry i mp l ica t ion . O n t h e o t he r h and , i fo ne w e r e s t udy ing ap opu la t ion k nown n o tt oc a r ry t h eg ene a noma ly f o r t h a la s sem ia , s u ch e nv i ronmen ta l i n f e r ence s c ou ld b e m ade . T hu s , k now ing t h e g ene t i c p r ed i spo s i t ion t o d i sea se o f i n d iv idua l s m ay h e lp i nt h e r e con s t ruc t ion o f t h e p a s t . T he n eed t o f o cu s o n t h e p o s s ib le e x i s t ence o f a n i n na te t e ndency t od eve lop o r a cqu i r e as p e c if ic d i sea se m u s t n o t b e u nde r -e s t ima ted , n o r s hou ld

t h e p o s s ib i l i ty o f r e t r iev ing t h i s i n fo rma t ion d i r ec t ly f r om s k e le ta l m a te r ia l . H ow , t h en , s hou ld w eo b ta in t h i sk i nd o fi n fo rma t ion?

S ource so fG ene t ic I nforma t ion o n D isea se S o f a r , t wo s o lu t ion s t o t h i s p rob lem h ave b een s ugge s t ed . O ne i s t o l o ok f o r s i gn s ( mo rpho log i ca l t r a i t s )o nt h es ke le ton w h ich s e em t ob e a s so c ia ted w i th s u scep t ib i l i ty t od i sea se ( s ee b e low ) ; t h e o t he r i st ot r y t od e te rm ine t h e r a c ia l a f f i l ia t ion o ft h e p opu la t ion s t ud ied . S ome t ime s , i n f a c t , t h e i d en t i f ica t ion o f t h e r a c ia l g r oup t o w h i ch t h ei n d iv idua l ss t ud ied b e long m ay i n d ica t e p a r t icu la rp r ed ispo s i t ion s t od i sea se . T h i si sd one o n t h eb a s i s t h a t s ome r a c ia l g roup s a r ek nown t o b e mo r e l i k e ly t o d eve lop c e r ta in d i sea s es t h an o t he r s . F i na l ly , a t h i rd s o lu t ion ( t he o ne s u gge s ted i nt h i s p ape r ) c on s ide r s t h e p o s s ib i l i ty o f o b ta in ing t h i s i n fo rma t ion f r om b i omo le cu le s w h ich m ay b e p r e sen t s t i l l i n t h e b one m a te r ia l i t s e l f . A t t h i s p o in t i tw ou ld s e em c onven ien t , s i nce g ene t ic s a nd r a ce a r e t e rms t h a t w i l l b e e ncoun te red f r equen t ly , t od ef ine w ha t i sm ean t b yr a ce i nt h ec u r ren tc on tex t . F o ro u rp u rpo se s ,a r a ce c an b e d e f ined a s a d i s t inc t g r oup o f i n d iv idua l s w h i ch d i s t ingu i she s i t se lf f r om o t he r p opu la t ion s b y a g ene p oo l i n w h ich g ene f r equenc ie s a r e c ha ra c te r i s t ic o f t h a t p opu la t ion a nd o f n o o t he r ( Thomp son a nd T homp son , 1 966 ) . I n o t he r w o rd s , r a ce s w i l l d i s t ingu i sh t h em se lve sa sd i f f e r ing i ns ome o ft h e g ene s t h ey c a r ry . T h i s i s ap u r e ly " g eno typ ic " d e f in i t ion , a s o ppo sed t o a" pheno typ i c " o ne . Whe r ea s t h e g eno type i s t h e g ene t i c c on s t i tu t ion i t s e l f , c ompo sed o f d i f f e r en t g ene s w h ich a r e i n t u rn " made o f ' DNA , t h ep heno type i st h ee n semb le o f e a s i ly s e en e x t e rna l c ha r ac te r i s t i c s . T he p heno typ e m ay r ef le c tc l o se ly t h e g eno typ e ,b u t i t m ay a l so b e t h e r e su l t o f " e nv i ronmen ta l " f a c to r s w h i ch i nf l uence a nd s omewha t a l te r t h e e xp r e s s ion o f t h e g eno type . S o t h e p heno type d o e sn o tn ece s sa r i ly e xh ib i tt h e g eno type ( Dav ie s , 1 986 ) . T he g eno type , o n t h e o t he rh and , w i thou t e x te rna l b i a se s , c ons i s t s o f t h e g ene t ic c onno ta t ion so fe a ch s i ng le p e r son . S o e ven i fi n s ome a u tho r s ' d e f in i t ion s p heno typ ic c ha r a c te r i s t i c s c ome i n to t h e d e s c r ip t ion o fr a ce , i ti ss a f e r , w hen t r y ing t od i f f e ren t ia te g r oup s a nd w hen s t udy ing p opu la t ion o r ig in s , m ovemen t s a nd m ix ing , t or e ly o n g ene t ic i n fo rma t ion w h ich m ay n o t b e i mmed ia te ly n o t iceab le m o rpho log i ca l ly b u t w h ich c ome s d i r e c t ly f r om t h e g ene s t h emse lve s . S uch i n fo rma t ion c an o n ly b eo b ta ined b ys t udy ing t h e g ene t i c m a te r ia l i t se l f ( DNA ) o r m o le cu le s c l o se ly r e la t ed t oi t ,a s w e w i l l s e el a t e r . T h i sd e f in i t ion a nd t h i se n t i r ei s sue a r e p a r t i cu la r ly r e levan t t o a r chaeo logy a nd a n th ropo logy . I nf a c t , w hen d ea l ing w i th h uman s k e le ta l m a t e r ia l , i n t h e p a s t g ene t ic a f f i l ia t ion s a mong i n d iv idua l s a nd t h e d e f in i t ion o f r a ce i n

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g ene ra l w e re g r ea t ly b a sed o n mo rpho log ica l c ha rac te r i s t ic s o ft h e s k e le ton . T he i n f e r en ce s o n r a c ia l a f f i l ia t ion i nt h i s c a se a r e n o t v e r y r e l iab le , d ue t ot h e f a c t t h a t , e xcep t f o r v e ry o bv iou s a nd m a jo r m o rpho log ica l d i f f e r ence s , i ti s n o t y e t k nown h ow ,i fa ta l l ,a nd t ow ha te x t en t ,g ene sa r e r e spon s ib le f o r t h e se c ha ra c te r i s t ic s . I fi tw e re p o s s ib le , t h e re fo r e , t o o b ta in " g ene t ic " i n fo rma t ion d i r ec t ly f r om DNA , o r f r om c l o se ly r e la ted m o le cu le s , t h en t h e d e f in i t ion o f g ene t ica l ly r e la ted g r oup sw ou ld b em o r er e l iab le . A f te r t h i s n e ce s sa ry d i g r e s s ion i t i s n ow p o s s ib le t or e tur n t ot h e m e thod s o fs t udy ing t h e p r ed i spo s i t ion o f i n d iv idua l s t o d i sea s e . C once rn ing t h e f i r s t m e thod , s t ud ie s o f c r an ia l m o rpho logy h ave b een p e r fo rmed i na na t temp tt o o b se rve a ny c o r r e la t ion w i th s pe c i f ic d i sea se s . A s t udy b y O l iv ie r e t a l ( 1 975 ) o n ag r oup o f n i ne teen th c en tu ry s k e le ton s f r om L i sbon p o in t s o u t t h a t t h e r e s e em s t o b e a n i n t e r e s t ing r e la t ionsh ip b e tween s ku l l f o rm a nd d i f f e r en t ia l m o r ta l i ty b y t u be r cu lo s i s , p l ague a nd s ma l lpox . S ub je c t s w ho h ad d i ed f r om t h e se d i sea se s h ad a l o nge r h ead a nd f a ce . T he p r a c t i ca l a pp l i ca t ion s o ft h i s s y s tem a r e v e ry l i m i ted , h oweve r . A s f a r a s t h e s e cond m e thod , ra c ia l a f f i l ia t ion , i s c once rned , v a r iou s s t ud ie s h ave b een p e r fo rmed o n t h e mo rpho log ica la nd m e t r ic c ha ra c t e r i s t ic so f t h ec r an ia o fd i f f e r en tr a ce sa nd o n h ow s u ch d a ta m ay h e lp d i f f e ren t ia t e b e tween g r oup s o f p opu la t ion s . T h i s , i n t u rn , c ou ld i n s ome i n s tances i n d i ca te w h i ch i n d iv idua l s a r e m o re l i k e ly t o c on t ra c t s pe c i f ic d i sea se s . B u t h ow r e l iab le i sa ny k i nd o fi n fo rma t ion o b ta ined f r om g r o s s mo rpho log ica l c ha ra c te r i s t ic s o f t h e c r an ium o ro fa ny o t he rp a r to ft h es k e le ton ? T he u s e o f o s teo log i ca l t r a i t s a s as ou r ce o f g ene t i c /d i sea se re la ted i n fo rma t ion ( w i th t h e e xcep t ion o f p a tho log i ca l l e s ion s , o f c ou r se ) c an b e m i s lead ing a nd i n conc lu s ive , t h e m a in a r gumen t a ga in s t t h i s p ro cedu r e b e ing t h a t i ti s n o t y e t k nown h ow m e t r ic o rm o rpho log ica l t r a i t s a r e i n he r i t ed n o rh ow t h ey d eve lop . S chw ide tzky ( 1 979 ) i sc o r r e c t i nh i s o b se rva t ion t h a t m any o f t h e se c ha rac te r i s t ic s a r e n o t a ge s tab le , h av e a c on t inuou s v a r iab i l i ty a nd t h a t t h e c ond i t ion s f o r t h e d eve lopmen t o f t h e se t r a i t s h a s y e t t o b e d i scove r ed . T he r e fo r e , t h ough t h ey m ay h ave s ome p oo r ly u nd e r s tood s i gn if i cance , t h e se t r a i t s c anno t p rov ide a r e l iab le s ou r ce o f g ene t ic i n fo rma t ion , d e sp i t e B e r ry ' s ( 1 979 ) a t temp t s t o m ake t h em s e em r e levan t . S ome r e sea r che r s , B u ik s t ra ( 1 976 ) f o r e xamp le , h ave l o oked a t o s teo log ica l t r a i t s a t ap opu la t ion l e ve l w i th t h e p r ec i se i n ten t ion o f i n teg ra t ing t h em w i th o t he r a r chaeo log ica l d a ta , t h u s p r e s en t ing a n e xce l len t e xamp le o f i n te r -d i sc ip l ina ry b l end ing w h ich s hou ld b e e mu la ted . E v en s o , o s teo log ica l t r a i t s a s as ou r ce o fg ene t ic i n fo rma t ion o fa ny s o r tm ay n o t b e a s r e l iab le a se xpe c ted . O the r s ou r ce s o f t h i sk i nd o fi n fo rma t ion m u s tb ep u r sued . Whe r e , t h en , s hou ld o ne t u rn f o r r e l iab le g ene t i c i n fo rma t ion o b ta ined d i r ec t ly f r om

s k e le ta l m a t e r ia l ? T he a n swe r l i e s i nb i o log ica l ly i n fo rma t ive m o lecu le st h a tm ay s u r v ive i na nc i en t b one , i n p a r t i cu la r DNA a nd c e r ta in t y pe s o f p r o te in s w h i ch a r e ad i r ec t " p roduc t " o fg ene s . I f i tw e re p o s s ib le t or e t r ieve r e levan t b i omo le cu le s f r om h uman o s seou s a r chaeo log ica l m a t e r ia l t h ey w ou ld c e r ta in ly p rov ide , i nt h eo ry , amuch m o re r e l iab le s ou r ce o f g ene t ic i n fo rma t ion . B e fo re d i s cu s s ing t h e p a r t icu la r t y pe s o f b i omo lecu le s w h ich c an g i ve u s t h i s k i nd o f i n fo rma t ion i t w ou ld b e s e n s ib le t oc on s ide r h ow f e a s ib le t h e i r e x t ra c t ion i sf r om a nc ien ts k e le ta l m a t e r ia l .

T he E x trac t ion o fI nforma t ive B iomo lecu les V e ry l i t t le i sk nown c once rn ing t h ed i agene s i s a nd f a t e i n g ene r a l o f o rgan ic m a te r ia l w i th in b one a f te r h und r ed s o r t h ou sand s o f y ea r s . S t ud ie s h av e d e s c r ibed h ow c e l l s d i e a nd h ow o r gan ic m a te r ia l d eg rade s , b u t l i t t le h a s b een s a id a bou t t h e i nf luen ce o f t i me , s o i l c ond i t ion s , t empe ra tu r e , a moun t o f o xygen , l e ach ing e f f e c t s o f w a t e r a nd s o f o r th o n b i o log ica l mo lecu le s . P ro te in s a nd o t he r b i omo lecu le s h ave b e en i s o la ted f r om a nc ien t s k e le ta l m a te r ia l . U n fo r tuna te ly , a sH edge s ( 1 978 ) h a s p o in t ed o u t , t h e c omponen t s o f b one t h a t s e em t os u rv ive i n g r ea t a bundan ce a r e t h e o ne s w h i ch a r e l e s s i n fo rma t ive a nd l e s s s u i tab le f o r a cqu i r ing i n fo rma t ion o n d i sea se s u s cep t ib i l i ty a nd g ene t i cs . I n f a c t , m o s t r e leva n t m o le cu le s a r e q u i t e s o lub le a nd p e rhap s m o r e e a s i ly l o s t t ot h e e f f e c t s o f l e a ch ing , a nd a r e a l so m o r e a c ce s s ib le t o a ta ck s f r om m ic ro -o rgan i sm s i n t h e g round w h ich m ay a l te r t h e i r n a tu r e . T h i s l a s t f a c to r s e em s t ob e o ne o ft h e m a in r e a son s w hy s t ud ie s o n t h e p o s s ib i l i ty o f r e t r iev ing m o lecu le s o f t h e ABO b l ood g r oup s ( ABO b l ood g roup a n t igen s ) a r e s o a mb iguou s a nd u n re l iab le . A s e cond p r ob lem i st h a t t h e o r gan ic r e s idue s w h ich d o p e r s i s t i n t i me w i l l c e r ta in ly b e p r e sen t i n v e ry s ma l l q uan t i t ie s . T h i s o b s ta c le m ay b e o ve r come b y t h e u s e o f mode rn i mmuno log ica l l a bo ra to ry t e chn ique sc apab le o fd e tec t ing mo le 9 le sp r e sen t i nn anog ram ( 1 0 -p )t op i cog ram ( 1 01 z )a moun t s . G iven t h e se d i f f icu l t ie s , i tm igh t s e em v e ry u n l ike ly t h a t o r gan i c m a te r ia l w i l l s u r v ive i na ny t y pe o fa nc ien tb one ( o ra r te f ac t ) b u t s up r i s ing ly , m any b i omo le cu le s s u ch a sc o l lagen , as ub s tance f o und i n m o s t t i s sue s , h ave b een i s o la t ed b y G u r t le r e ta l( 1 98 1 ) . L oy ( 1 983 ) h a s d i s cove r ed s p e c i e s spe c i f ic h aemog lob in c r y s ta l s f r om 2 000 y ea r o l d b l ood s ta ined a r row h ead s a nd h a s a l so i d en t i f ied h uman b l ood f r om p r eh i s to r ic r o ck p a in t ing s ( Loy e t a l , 1 990 ) . U s ing r a d io i mmuno log i ca l a s say s , L owen s te in ( 1 980 , 1 981 , 1 985 ) h a s d e te c ted a l bum in ( a p ro t e in f o und i n t h eb l ood ) i n1 . 9 m i l l ion y ea ro l d f o s s i l s . H e h a s a l so f o und s pe c ie s -spe c i f ic p ro te in s i n N eande r tha l s ,H omo e r ec tu sa nd A u s t ra lop i thecus r obu s tu s ( 1 981 ) . A scenz i e t a l ( 1 985 ) h ave d e t e c ted i mmuno log i ca l ly a l pha a nd b e ta

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t h a t c once rn ing s ke le ta l mo rpho logy a nd s ke le ta l l e s ions . T he s e ar ch f o r a ny k ind o f b iomo le cu le i mp l ie s l o ok ing f o rs ome th ing o ne c anno ta c tua l ly s e e , a nd t h e re fo re i t c on s is t s , f i r s t o f a l l , o f e x t ra c t ing f r om t h e b one t h e s ub s tance o ne h opes i s p r e sen t a nd t h en u s ing a n a dequa te t e s t t o d e te c ti t s p r e sence . T he t e s t s m ay v a ry a cco rd ing t ot h e mo lecu le w h ich i sb e ing s ough t , a nd m ay b e mo re o r l e s s c omp lex . B u t t h e l o g ic b eh ind t h is k i nd o f r e sea r ch i s , g ene ra l ly s peak ing , t h e s ame r e ga rd le s s o fw ha t o ne i ss eek ing . T hey a r e t e s t s , o ra s say s , w h ich h ave b e en d eve loped i nt h e f i e ld o fb i omed ica l s c iences . A s a n e xamp le o ne c an c on s ide r t r y ing t o e x t ra c t af a i r ly s imp le mo le cu le : t h e b lood p ro te in a l bum in . T h is p ro cedu re c on s is t s , f i r s t o f a l l , o f t a k ing p e rhaps 1 0g .o fb one f r om t h e v e r teb rae , f emo ra l h ead o r , p e rhap s , t h e i l i ac c r e s t . T h is b one s hou ld b e c r ushed a nd t h en , f o r ap e r iod o f t ime , p u t i n to a s o lu t ion w h ich s hou ld h e lp e x t rac t t h e a l bum in f r om t h e b one . A f te r t h i s , t h e s o lu t ion i nw h ich t h e b one p owde r w a s i mme r sed i s c o l le c ted . H ope fu l ly i tw i l l c on ta in s ome a l bum in . T h is a moun to fl i qu id ( t he e x t ra c t ) i st h en c oncen t ra ted t o af ew m i l l i l i t res , f o r e xamp le , a nd t e s ted . F ind ing t h e a pp rop r ia te t e s t w i th w h ich t od e tec t t h e p r e sence o f a l bum in i nt h e e x t rac t i sa no the r c r uc ia l s t ep . T he t e chn ique mus t b e b o th v e ry s e ns i t ive , i no rde r t od e tec t v e ry s ma l l q uan t i t ie s o f a l bum in , a nd v e ry s pec if ic , s o t h a t a no ther mo le cu le , c hem ica l ly s im i la r t oa l bum in , w i l l n o t b e e r roneou s ly p i cked u p a s a l bum in . A n e xamp le o f s u ch at e s t i st h e t e chn ique k nown a s E L ISA ( e nzyme l inked i mmuno so rben t a s say ) . T h is s y s t em u se s mo lecu le s c a l led a n t ibod ie s , w h ich w i l lr e cogn ize a nd " s t ick t o " o n ly as pec if ic mo lecu le . D if f e r en t a n t ibod ies c an b e m ade t o r e cogn ize d i f f e r en t mo lecu le s . S o i f as o lu t ion o f a n t ibod ie s w h ich w i l l r e cogn ize o n ly a l bum in i s m ixed w i th t h e e x t r ac t ( t h is i s u sua l ly d one i n v e ry s ma l l w e l l s o f ap l a s t ic p l a te ) , a nd i fa ny a l bum in i sp r e sen t i nt h e e x t r ac t , t h ea n t ibody w i l l " a t tach " t o i t . T h is c anno t b e v i sua l i sed i mmed ia te ly , s o as e cond a n t ibody w h ich w i l l r e cogn ize t h e f i r s t o ne h a s t o b e u sed . T h is s e cond a n t ibody a l so c anno t b e " s een " , b u t i th a s a t tached t oi tam o le cu le w h ich w i l l i nduce at h ird s ub s tance f i na l ly t o c ause a f ou r th a dded s ub s tance t ot u rn ac e r ta in c o lou r . S o t h ep ro ce ss o f d e te c t ing a l bum in c on s is t s o f a dd ing o ne s ub s tance a f te r a no the r , e ach o ne o f w h ich w i l l a t tach t o t h e p r ev ious o ne u n t i l , f i na l ly , t h e a dd i t ion o f ac o lour ing a gen t w i l l a l low u s t os ay w he the ra l bum in i sp r e sen t i nt h a t p a r t icu la rb one e x t ra c t o r n o t . I ti so bv iou s t h a t t h e l o g ic t o f o l low i np e r fo rm ing t h e se e xpe r imen t s , a nd t h e s e r ie so fd i f f icu l t ie so ne h a s t oc onf ron t ,a r e n ove l t ot h ef i e ld o fa r chaeo logy . T he r ema in ing p a rag r aphs o f t h i s c hap te r c oncer nt h ea bove -men t ioned HLA a n t igen s . T he c ho ice o ff o cu s ing o n t h e se p ro te in s w a s d i c ta ted b y t h e i r c apa c i ty , p e rhap s mo re t h an t h a t o f a ny o t he r b i omo le cu le e xcep t f o r DNA , t o s upp ly

h aemog lob in c ha ins i nb one u p t o4 500 y ea r s o ld , a nd R o thsch i ld a nd T u rnbu l l ( 1987 ) s e em t oh ave d e te c ted , v i a i mmunoh is to log ica l t e chn ique s , a n t igens o f t h e T reponema b ac i l lum i nl e s ions o f aP le i s tocene b ea r . A ga in w i th i mmuno log ica l m e thod s U lr ich ( 1987 ) h a s e x t rac ted o s teoca lc in ( ap ro te in p re sen ti nt h em ine ra lp ha se o fb one ) i n a n ima l f o s s i ls f r om 1 2 ,000 t o 1 3 m i l l ion y ea r s o ld . H e a l so s ub jec ted b one t o v ery h i gh t empe ra tu res ,a sd i dL owens te in w hen l o ok ing f o r a l bum in , a nd f ound t h a t t h e a n t igen ic a c t iv i ty o f t h e mo lecu le , i t s d i s t ingu ish ing c hem ica l c ha ra c te r i s t ics i no the rw o rd s ,h ad b een r e ta ined . DNA h a s b een d e tec ted ( J one s , 1 986 ; P aabo , 1 986 ) i n m umm if ied t i s sue a nd r e cen t ly ( Hage lbe rg e t a l , 1 989 ) i n h uman s ke le ta l r ema ins . When l o ok ing f o r g ene t ic i n fo rma t ion c once rn ing c ha r ac te r is t ics s uch a s r a ce a nd s u scep t ib i l i ty t o d i sea se , t h e a va i lab i l i ty o f t h e DNA mo lecu le i t se lf , o r f r agmen t s o f i t , w ou ld s upp ly t h e mo s t d i re c t g ene t ic i n fo rma t ion a va i lab le a nd w ou ld i nt h eo ry b e t h e mo s tr e l iab le s ou rce o f s uch i n fo rma t ion a mong m o s t o t he r b i omo lecu le s . B u t , a s P aabo ( 1986 ) h a s p o in ted o u t , a l though DNA s e ems t ob e r e s i s tan t t oh a r sh a gen t s , t h e re a ppea r t o b e p rob lems c once rn ing t h e v e ry l ow y i e ld o b ta ined w hen a t temp t ing t o c l one a nc ien t DNA . V enanz i a nd R o l lo ( 1990 ) h ave e ven s ugge s ted t h a t t h e n uc le ic a c id s ( c omponen t s o f DNA ) i n a nc ien t m umm ies a r e m a in ly o f RNA ( a no the r t y pe o fg ene t ic m a te r ia l , d i f f e ren t f r om DNA ) . A l though H age lbe rg e ta l ( 1 989 ) h ave s ucceeded i n i s o la t ing DNA f r om s ke le ta l ma te r ia l , f u r the rr e sea r ch mus tb ed one i n o rde rt oc onf i rm t h is ,a nd e spe c ia l ly t oc on t ro l f o r t h e p o s s ib i l i ty o fc on tam ina t ion f r om f r esh DNA , s e e ing t h a t t h e e xpe r imen t e mp loys a n e x t r eme ly s e ns i t ive t e chn ique . N one the le s s , i f DNA d oe s s u rv ive i n a nc ien t m a te r ia l , t h e i mp l ica t ions f o r t h e f i e ld s o fa r chaeo logy a nd a n th ropo logy w ou ld b e v e ry s i gn if ican t . A no the r g roup o f b i omo le cu le s w h ich w ou ld b e e x t r eme ly h e lp fu l i nt h ese k i nd s o f i n ves t iga t ions ( a nd w ou ld b e a c l o se s e cond t o DNA i n t h e t y pe o f g ene t ic i n fo rma t ion t h ey c an p rov ide ) i st h a t o ft h e HLA a n t igens , w h ich w i l l b e c ons ide red i n d e ta i l f u r the r o n . A l though t h e e x t r ac t ion o f t h is " g lycop ro te in " ( i . e . HLA ) c u r ren t ly p r e sen t s d i f f icu l t ies e ven w he r e r e la t ive ly f r esh m a te r ia l i s c once rned , t h e p o s s ib i l i ty o f i t s s u rv iva l i n a nc ien tm a te r ia ls hou ld b ec ons ide red s e r iou s ly . G ene ra l ly s peak ing , t h e c ause s a nd m echan isms o f s u rv iva l o f t h e se mo lecu le s a r e n o t k nown . A p a r t ia l e xp lana t ion m ay b e p rov ided b y t h e p a r t icu la r b io chem ica l c ha rac te r is t ics o f t h e mo le cu le s o r , p e rhap s , b y a c e r ta in d eg r ee o f p ro tec t ion g i ven b y t h e a pa t i te ( i . e . m ine ra l ) m a t r ix o r t h e t r abe cu la rs t ruc tu r e i n c e r ta in p a r t s o f t h e b one i t se l f . I n c onc lus ion , i t h a s b een p roven t h a t v a r iou s o rgan ic mo le cu le s d os u rv ive i nh uman o s se iou sm a te r ia l . I ti si mpo r tan tt ou nde r s tand h ow d i f f e ren ta nd h ow d e l ica te t h i s k i nd o fr e sea rch i sc ompa r ed t o

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" o rde r ed " b y an umbe r o f d i f f e ren t g ene t ic l o c i . T hey h ave b een s e pa ra ted i n to t h ree c l a s se s : C la s s I ,C la s s Ia nd C la s s I I ( S ch ieb l , 1 986 ) . F o cu s ing o n C la s s Ia n t igen s , f o r e xamp le , o ne c an s e e t h a t t h ey a r e p roduc t s o fl o c i A , Ba nd C a nd t h a t t h ey a r e e xp re s sed o n t h e s u r f a ce o f a l l n uc lea ted c e l l s a nd p l a te le t s ( Bende r , 1 984 ) ( b e cau se o f t h e i r v a s t d i s t r ibu t ion , t h ey s e em t o f u nc t ion a s m a rke r s o f s e lf r ecogn i t ion ; i n f a c t , o n ly i n t h e c on t ex t o f s e l f recogn i t ion m ay o r gan i sms r e cogn ize n on se l f a nd t a ke i mmuno log ica l a c t ion a ga in s t i t ) . O ne l o cu s c an h ou se d i f f e r en t v a r ian t s o f ag ene w h ich a r e c a l led a le le s . T he se a l le le s w i l l c ode f o r d i f f e ren t " va r ie t ie s " o f t h e s ame p ro te in ( e .g . HLA a n t igen ) ( F igu r e 5 . 3 ) . S o l o cu s A , f o r e xamp le ,c an c ode f o r u p t o2 1 d i f f e ren t t y pe s o f HLA -A a n t igen s ( Thom son , 1 986 ) ,o fw h ich e a ch i n d iv idua l c an o n ly d i sp lay t wo . T he a l le le s a r e a s s igned n umb e r s p re ce ed ed b y t h e l e t te r o f t h e r e spe c t ive g ene l o cu s . T ab le 1g i ve s as umma ry o ft h ed i f f e r en tk i nd so f HLA a n t igen s . E a ch i n d iv idua l c an e xp re s s n o mo r e t h an t wo a n t igen s f o r e a ch l o cu s . T he HLA a n t igen c omb ina t ion f o r C la s s I , I , I I a n t igens , d e t e rm ined b y t h e g ene s o f o ne c h romo some , i s t h e h ap lo type , o fw h ich e ve ry i n d iv idua l h a s t wo , o ne o fe a ch h omo logou s c h romo some . H uman s , i nf a c t , h ave 4 6 c h romo some s ( t he s t ruc tu r e s i n to w h ich g ene t ic m a te r ia l i s " r o l led " u p ) w h i ch c on s i s t o f 2 3 s e t s o f t wo h omo logou s c h romo some s , i . e . t wo s t ruc tu r e s c on ta in ing t h e s ame k i nd o f i n fo rma t ion , o ne s e t c om ing f r om t h e m o the r , t h e o t he r f r om t h e f a the r . T he h ap lo typ e i sp a s sed o n a s au n i t . T hu s , e a ch o f t h e t wo p a ren t s w i l l o f f e r t wo h ap lo type s , b u t o n ly o ne f r om e a ch p a r en t w i l l b e r a ndom ly a c cep t ed , s o t h a t f r om t wo p a ren t s , o n ly f o u r h ap lo typ e c omb ina t ion s a r e p o s s ib le ( F igu r e 5 . 4 ) . A n e xamp le o f a n i n d iv idua l ' s HLA t y p e , c on s ide r ing o n ly t h e Aa nd B l o c i f o r t h e s a ke o f s imp l ic i ty , c ou ld b e : A l , A ll , B 8 , B 35 o r A l , A 3 , B 7 , B 8 o rm any o t he rc omb ina t ions o f Aa nd B .

g ene t ic i n fo rma t ion . T he d ec i s ion t o f o cu s o n HLA i n s tead o f DNA w a sm ade p a r t ly b e cau se o f t h ep e r sona le xpe r ience t h ea u tho rh a si ns t udy ing a nd a t temp t ing t o e x t ra c t HLA f r om a nc ien t m a te r ia l , p a r t ly b e cau se i tc ou ld p rove t ob e a n a l te rna t ive m e thod ( w i th s ome a dvan tage s a nd , e qua l ly , s ome d i sadvan tage s o ve r DNA , b u t w h ich h a s n o t r e ce ived a s m uch a ten t ion ) f o r o b ta in ing g ene t ic i n fo rma t ion .

HLA An t igens a s aS ource o fG ene t ic I nforma t ion I fw e a r e t ol o ok f o r g ene t ic i n fo rma t ion , w e m u s t c hoo se a mo le cu le w h i ch w i l l r e f le c t b e s t t h e i n fo rma t ion c on ta ined i n o u r g ene s . O ne c and ida te i st h ec omp lex b i omo le cu le c a l led HLA a n t igen ( Human L euko cy te A n t igen ) , w h i ch i s a ncho red t ot h e s u r f a ce o fm o s t c e l l s i no u rb ody a nd w h ich r e gu la te s o u r i mmune r e spon se . T he e n t i re HLA s y s tem , w ho se c oun te rpa r t i n o u r g ene s i s c a l led t h e Ma jo r H i s to compa t ib i l i ty C omp lex , i st h e mo s t r ema rkab le g ene t ic m a rke r . I tl e nd s i t se lf f a vou rab ly t os t ud ie s , n o t o n ly o f s e lec t ion , h uman e vo lu t ion a nd g ene t i c s ( Bende r , 1 984 ) ( f o r p a te rn i ty t e s t s a nd g ene t ic c ha r ac te r isa t ion o f d i f f e ren t h uman p opu la t ion s , f o r e xamp le ) b u t i ta l so , a nd e spe c ia l ly f o r o u r p u rpo se s , l e nd s i t se l f t o d i sea se a s so c ia t ion s t ud ie s . a )B io logy o f HLA a n t igens T he mo re c omp lex d e ta i l so ft h i sm o lecu le a r e b eyond t h e s cope o f t h e c hap t e r . N one the le s s , a b r ie fd e sc r ip t ion o fi t s s t ruc tu re a nd f u nc t ion m ay p e rhap s h e lp t o u nde r s tand i t s i mpo r tan ce . T he Ma jo r H i s to compa t ib i l i ty C omp lex ( MHC ) i sa c omp lex o fg ene s w h i ch c ode s f o rt h ep r oduc t ion , b y t h e c e l l , o f HLA a n t igen s . I n m an , HLA a n t igen s w e r e c a l led H uman L euko cy t e A n t igens b ecau se t h ey w e re f i r s t s e en o n l ympho cy t e s , a t y pe o f w h i te b l ood c e l l . T he se m o le cu le s a r e p r o t e in s , mo re p r ec i se ly g l y cop ro te in s , w h i ch a r e e xp r e s sed a nd e mbedded w i th in t h es u r f a ce o ft h e c e l l s o fa l l h i ghe r v e r t eb r a te s a nd w ho se f u nc t ion i s t o r e gu la te t h e i mmune r e spon se w h i ch i s c r uc ia l t o t h e s u rv iva l o f i n d iv idua l s ( Bende r , 1 984 )( F igu re 5 . 2 ) . B r ief ly , g ene s ( DNA ) p rov ide i n fo rma t ion w h ich t h e c e l l m ay t r an s la t e i n to p ro te in s . T he i n fo rma t ion s e n t b y t h e g ene s o r de r s t h e c e l l t o s t r ing t o ge the r a s e r ie s o f " bu i ld ing b l o ck " m o lecu le s c a l led a m inoa c id s t op roduce ap ro t e in m o lecu le s uch a s a n HLA a n t igen . D i f f e r en t c omb ina t ion s o f al i m i ted v a r ie ty o f a m inoa c id s a nd d i f f e ren t l e ng th s o f t h e se s t r ing s w i l l a low f o r d i f f e ren t k i nd s o f p ro t e in s ( t he re a r e m any m o re c omp l ica ted f u nc t ion s i n vo lved i n t h i s i n s tance , b u t f o r t h e p u rpo se o f t h i s c hap t e r , t h i s d e f in i t ion w i l ls u f f ice ) . Al o cu s i s ap o s i t ion o n t h e g ene t ic m a t e r ia l w h i ch h ou ses o ne g ene . HLA a n t igen s c an b e

b ) HLA a n t igens a si nd ica tors o fr ace T he f r equency o f HLA t y pe s , a nd o f s pec i f ic c omb ina t ion s o f t h e se , i s i mpo r tan t i n c ha ra c te r i s ing p opu la t ion g r oup s a nd r a c ia l a f f i l ia t ion . T ab le 5 . 2A s how s al i s t o f t h e se d i f f e r ence s , a s d o e s T ab le 5 . 2B f o r p opu la t ion s w i th in a nd c l o se t oE u rope . F o r e xamp le , HLA A l , A3 , B8 a r e f o und m a in ly a mong C auca s ian s ; A24 , AW31 , B40 a mong o r ien ta l s a nd AW30 , B17 a nd BW42 a mong N eg ro e s ( Bend e r , 1 984 ) . I ti sk nown t h a td i f f e ren ta r ea so f E u rop e d i f f e r i n HLA f r equenc ie s , e spe c ia l ly N o r the rn a nd We s t e rn E u rope f r om c en t ra l a nd E a s t e r nE u rop e( Bodme ra nd B odme r , 1 978 ) .

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c ) HLA a n t igens a sp os s ib le i nd ica tor so f d i sease

c ance l lou s b one a nd f r om t h e f r agmen t s o f c e l l m emb rane i nw h ich i ti sp o s s ib ly s t i l l e mbedded . Mo re r e sea r ch n eed s t ob e d one b e fo re a ny th ing m ay b e s a id c once rn ing t h e f e a s ib i l i ty o f e x t ra c t ing t h e se mo le cu le s .

B e s ide s h e lp ing u s t o d e te rm ine t h e r a c ia l a f f i l ia t ion o f i nd iv idua l s , a nd t h e re fo re h e lp ing i n d i rec t ly t o p rov ide i n fo rma t ion o n p r ed i spo s i t ion t o d i sea se , t h e se a n t igen s c an b e , t h emse lves , d i r ec t ly a s soc ia ted w i th p a r t icu la r d i sea ses ( Tab le 5 . 3 ) . A s so c ia t ion i n t h i s c a se m eans , i n mode rn m ed ica l t e rms , t h a t t h e p a t ien t p opu la t ion ( Thomson , 1 986 ) h a s as i gn if ican t ly i n cr ea sed f r equency o f ap a r t icu la r I I LA a n t igen c ompa red t o ac on t ro l p opu la t ion . S ome o ft h ese a s so c ia t ion s a r e n o t v e ry s t rong . O n t h e o t he r h and , s ome a r e q u i te m a rked , s u ch a s t h a t b e tween HLA -B27 a nd a nky lo s ing s pondy l i t is . R esea r ch h a sb een c a r r ied o u to nt h e se a n t igen s t o e l uc ida te t h e i r p a r t icu la r r e la t ionsh ip w i th i n fec t ious d i sea se s s uch a s s ma l lpox , m a la r ia , c ho le ra , t u be rcu lo s is ( Bodme ra nd B odme r , 1 978 ) a nd l e p ro sy ( d e V r ie s e ta l , 1 976 ) . I ti sc l ea r , t h e r efo r e , t h a t t h e p r e sence o f c e r ta in HLA a n t igens m ay i n d ica te s u scep t ib i l i ty t o c e r ta in d i sea se s . T he s us cep t ib i l i ty a s soc ia t ed w i th t h e p r esence o f t h e se p a r t i cu la r a n t igens m ay b e a d i re c t e f f ec t o f t h e HLA a n t igen , b u t i n mo s t c a se s t h e HLA a n t igen i ss imp ly ama rke rc l o se ly l i nked t o al o cus w h ich i sc au sa l ly r e la ted t ot h e d i sea se . T he p o in t r ema ins t h a t t h e se mo lecu le s a r e p o ten t ia l ma rke r s f o r d i sea se . T he se a n t igen s m ay e ven g i ve u s s ome i n s igh t i n to w ha ti lne sse s t h ese p opu la t ion s m ay h ave f a ced , e ven i ft h ey d i d n o t l e ave t h e i r m a rk o n b one , w h ich h a s b een t h e o n ly s ou r ce o f e v idence f o r d i sea se a va i lab le t ot h ep a laeopa tho log is tu pu n t i ln ow .

C onc lus ion Ih ope I h ave s ucceeded i n s t re s s ing t h e i mpo r tance , a s w e l l a s t h e d i f f icu l t ie s a nd t h e n eed f o r f u r the r r e sea r ch , i n a cqu i r ing g ene t ic i n fo rma t ion o n r a ce a nd h ea l th d i rec t ly f r om b one i n t h e f o rm o f b i o log ica l mo lecu le s , r a ther t h an f r om l e s s r e l iab le s ou r ces s uch a st h e s t udy o ft h e g ro s s mo rpho logy o f t h e s ke le ton o r o f b lood g roup s o r HLA a n t igen d i s t r ibu t ion a mong m ode rn p opu la t ions . I n f a c t , b iochem ica l a nd s e ro log ica l s t ud ie s c on ce rn ing g ene t ic t r a i t s a nd v a r iab i l i ty h ave b een p e r formed o n mode rn p opu la t ions i nt h e a t temp t t os hed s ome l i gh t o n h uman g ene t ic p opu la t ion p a t te rn s o ft h e p a s t , b y e x t rapo la t ing m odem d a ta t oa n a nc ien t c on tex t . H oweve r , b e cau se i ti sn o t p o s s ib le t oc on t ro l f o r t h e g r ea t n umbe r o f f a c to r s a nd v a r iab le s w h ich o ve r t h e m i l lenn ia c ou ld h ave a f f ec ted t h e f i na l g ene t ic m ake -up o f mode rn p opu la t ions , " gene t ic" s t ud ie s p e r fo rmed d i rec t ly o n a nc ien t b one a r e i n t h eo ry mo re r e l iab le . We s hou ld t h e r e fo re d i rec t o u r a t ten t ion t o d e te rm in ing w ha t h appens t ot h e se mo lecu le s a nd h ow t h ey m ay b e r e t r ieved , t o d ec iphe r ing t h e i n fo rma t ion o n i n na te s u scep t ib i l i ty t od i sea se ( o r a ny o t he r g ene t ic k i nd o f i n fo rma t ion w e m ay n eed ) , a nd t h en t o i n teg ra t ing t h is e s sen t ia l b a ckg round i n fo rma t ion i n to t h e r e s t o f t h e a r chaeo log ica lp i c tu re ( F igu r e 5 . 5 ) . A b r ief m en t ion s hou ld a l so b e m ade c once rn ing t h e i mpo r tance , i n t h e c on tex t o f d i sea se , o f l o ok ing f o r b i omo lecu les t h a t a r e n o t n ece s sa r i ly h uman ; f o r e xamp le , m o lecu le s c ha r ac te r is t ic o f s pe c i f ic m ic ro -o rgan isms t h a t a f f ec t m an a nd w h ich m ay s t i l l b e p r esen t i n l e s ion s o n t h e s k e le ton o ft h e i nd iv idua l . I no t he r w o rd s , l e s ion s c au sed b y i n f ec t ious d i sea se s s uch a s t u be r cu lo s i s o r s yph i l i s m ay s t i l l c on ta in d i agno s t ic b i omo lecu le s o f t h e s pec i f ic b a c te r ia l a gen t s r e spons ib le f o r t h e o nse t o f t h e d i sea se . T hese m ay a low u s t o m ake a mo re a ccu r a te d i agno s is . F ina l ly , t h e p o in t Iw ou ld l i ke t o c onvey i s t h a t n o m a t te r w h ich p a r t icu la r b iomo lecu le w e m igh t c ons ide r r e cove r ing f r om s ke le ta l m a te r ia l , b e i tDNA , HLA a n t igen s o rb lood p ro te ins , t h ese g ene t ica l ly i n fo rma t ive o r d i sea se -a s soc ia ted m o lecu les d o e x i s t a nd t h e re i s ar e a l p o ss ib i l i ty t h a t t h ey m ay p e r s is t i n t ime . T he re fo re mo re e f fo r t s hou ld b e s pen t i nt r y ing t ou nde r s tand t h e p ro ce s se s o f d eg rada t ion o f t h ese mo lecu les a nd t h e c ond i t ion s o f t h e i r s u rv iva l . T he i r r e t r ieva l a nd e x t rac t ion f r om h uman s ke le ta l m a te r ia l c an g i ve u s s ome c l ue s c once rn ing t h e g ene t ic ( i nna te ) s u s cep t ib i l i ty t od i sea se . T h is w ou ld , i n t u rn , a l low as ounde r b a se f o r t h e r e con s t ruc t ion

d ) HLA a n t igens : w ork t od a te A l i t e ra tu re s ea r ch h a s f a i led t o s how a ny t h o rough i n ves t iga t ion o n HLA a n t igen s i nb one s a pa r t f r om t h e a t temp t m ade b y H edge s a nd Wa l la ce C a rm ichae l ( 1 978 ) i nw h ich t h e a n t igen s w e re n o tf ound . E x t r ac t ion o f HLA a n t igens f r om S axon a nd Med ieva l h uman b one h a s b een a t temp ted a t t h e R eg iona l B lood T ran sfu s ion C en t re i nS hef f ie ld ( Ca t taneo , 1 988 ) . T he m e thod u sed f o r t e s t ing e x t ra c t s m ade f r om f r e sh a nd a nc ien t h uman b one ( a s d e sc r ibed p r ev iou s ly i n t h is a r t ic le ) w e r e s im i la r t o t h o se u sed i n t i s sue t y p ing mode rn i n d iv idua ls ( i . e . d e te rm in ing t h e i r HLA c ha ra c te r i s t ics , w i th av i ew t ot r ansp lan t so f p a r t icu la r o rgans o r t i s sue s , f o r e xamp le ) . T h i s t e s t , c a l led l ymphocy to tox ic t e s t , w a s a l so e mp loyed b y S ta s tny ( 1973 ) t o t e s t mumm if ied s o f tt i s sue f o rt h ep re sence o f HLA a n t igen s . T he d e ta i l so ft h e e xpe r imen t s p e r fo rmed a tt h e RBTS a r e b eyond t h e s cope o f t h i s c hap te r . T he e x t r ac t ion m e thod s e mp loyed s ucceeded i n r e t r iev ing a l bum in ,b u tp rob lem s a r o se w i th HLA . T he d i f f icu l ty i n e x t rac t ing t h i s m o le cu le , i f p re sen t , f r om a nc ien t o r modem b one l i e s e spec ia l ly i nt h e d oub le n a tu re o f t h e e x t rac t ion . I t mu s t b e r e t r ieved b o th f r om t h e r e ce s se s o ft h e

4 4

L owen s te in , J .M . ( 1 985 ) Mo lecu la r a pp roache s t ot h e i d en t if ica t ion o fs p ec ie s . Am .S c i . 7 3 :541-47 . L oy , T .H . ( 1 983 ) . P r eh i s to r ic b l ood r e s idue s : d e tec t ion o n t o o l s u r f a ce s o fs pec ie s o r ig in . S c ience 2 20 :1269 -7 1 . L oy , T .H . , J one s ,R . , N e l son , D .E . , Meehan , B . , V oge l , J . , S ou thon , J .& C o sg rove , R . ( 1 990 ) . A cce le ra to r r a d ioca rbon d a t ing o fh uman b l ood i np i gmen t s f r om L a te P l e i s tocene a r t s i te s i n A u s t ra l ia . A n t iqu i ty 6 4 :110 16 . O l iv ie r , G . , d e C a s t ro , M .E . &A lmed ia ( 1 975 ) . S ku l l f o rm a nd d i f f e ren t ia l mo r ta l i ty b y t u be rcu lo s i s . J . H um .E vo l .4 : 491-95 . O r tne r , D . J . a nd P u t scha r , W .G .J . ( 1985 ) . I d en t i f ica t ion o fP a tho log ica l C ond i t ions i nH uman S ke l e ta l R ema ins . Wa sh ing ton : Sm i th son ian I n s t i tu te P r e s s . P aabo , S .( 1 986 ) . Mo lecu la r g ene t ic i n ve s t iga t ion s o f a nc ien t h uman r ema in s . C o ld S pr ing Ha rbo r S ympo s ia o nQ uan t i ta t ive B io logy 2 1 :441-46 . R o th sch i ld , B .M . a nd T u rnbu l l , W . ( 1987 ) . T reponema l i n f ec t ion i n aP l e i s tocene b ea r . Na ture 3 29 :61-2 . S ch ieb l , B . ( 1 986 ) . HLA t y p ing . P rob lem s a nd s o lu t ion s . B io te s tM i t t . ( s pec ia le d i t ion ) 7 . Mun ich : B io t e s t AG . S chw ide tzky , I .( 1979 ) . P a laeopopu la t ion g e n e t ic s . J . H um .E vo l .8 : 669 -77 . S t a s tny , P .( 1 973 ) . HLA a n t igen s i nm umm i f ied P re C o lumb ian t i s sue s . S c ience 1 83 :864 -66 . S t e inbock , R .T . ( 1976 ) . P a leopa tho log ica l D iagno s i s a nd I n te rp re ta t ion . S p r ingf ie ld : C ha r le sC .T homa s . S ve jgaa rd , A . a nd R yd e r , L .P . ( 1 977 ) . A ssoc ia t ion s b e tween HLA a nd d i sea se . I n HLA a nd D i sease e d s . J . D au s se t a nd A . S ve jgaa rd . p p . 4 6 -53 . C openhagen : Munk sgaa rd . T homp son , J . S . a nd T homp son , M .W . ( 1 966 ) . G ene t ic si n Med ic ine . P h i lade lph ia : W .B . S aund e r s . T hom son , G . ( 1986 ) . H uman HLA g ene t ic s a nd d i sea se a s soc ia t ion s . I n G ene t ic s a nd Mo lecu la r I mmuno logy e d . D .M . We i r H andbook o f E xpe r imen ta l I mmuno logy 3 . p p . 1 102 . 12 O xfo rd : B lackwe l lS c ien t if ic P ub l ica t ion s . U l r ich , M .M .W . , P e r izon iu s , W .R .K . , S poo r , C .F . , S andb e rg , F . & V e rmee r , C . ( 1 987 ) . E x t rac t ion o f o s teoca lc in f r om f o s s i l b one s a nd t e e th . B io ch em . B iophys .R es .C omm . / 49 :713 19 . V enanz i , F .M . a nd R o l lo , F . ( 1 990 ) . Mummy RNA l a s t sl o nge r . N a ture 3 43 :25 .

o f t h e h ea l th s t a tu s o f ap opu la t ion a nd , t o ge the r w i th o the ra r chaeo log i ca ld a ta ,w ou ld h e lp d r aw a mo re a ccu ra t e p i c tu r e o ft h e l i f e s ty le a nd c u l tu ra l c hange so fp a s tp opu la t ion s .

R e f erence s A lbe r t s , B . , B ray , D . , L ew i s , G . , R a f f , M . , R obe r t s ,K . & Wa t son , J .D . ( 1983 ) . T he C e l l . N ew Y o rk : G a r land . A nge l , J . L . ( 1967 ) . P o ro t ic h ype ro s to s i s o r o s t eopo ro s i s s ymme t r ica . I n D i sea ses i nA n t iqu i ty e d s . D . B ro thwe l l a nd A . S and i son . S p r ingf ie ld : C ha r le sC .T homa s . A scenz i , A . , B runo r i , M . , C it ro , G . &Z i to , R . ( 1985 ) . I mmuno log ica l d e tec t ion o fh aemog lob in i nb one s o f a nc ien tR oman t ime s a nd o fi r on a nd e neo l i th ic a ge s . P roc .N a t .A cad .S c i .8 2 :7170 -72 . B ende r ,K . ( 1984 ) . HLA s y s t em . B io .B u l l .2 : 61131 . B e r ry , R .J . ( 1979 ) . S ec t ion I . G ene s a nd s k e le ton s a nc ien ta nd m odem . J .H um .E vo l .8 : 661 -67 . B odme r , W .F . a nd B odme r ,J . G . ( 1978 ) . E vo lu t ion a nd f unc t ion o f t h e HLA s y s tem . I nT he HLA S y s tem . e d W .F . B odme r , p p . 3 09 -24 . B r i t . Med . B u l l . 3 4 :213 3 24 . B u ik s t ra , J .( 1976 ) . H opew e l l i n t h e L owe r I l ino i s V a l ley . N o r thwe s te rn U n ive r s i ty A rchaeo logy P rog ram S c ien t i f ic P ape rs ,N o .2 . C a t taneo , C . ( 1 988 ) . P a laeo in zmuno logy : t h e I mpo r tance o f HLA A n t igen s i nA rchaeo logy a nd t h e F eas ib i l i ty o f t h e i r R e t r ie va l f rom A nc ien t H uman S ke le ta l Ma te r ia l . U npub l i shed M .A . t h e s i s , U n ive r s i ty o fS hef f ie ld . D au s se t , J . a nd C o lomban i , J . ( 1 973 ) . e d s . H is tocompa t ib i l i ty T e s t ing R epo r t . C op enhagen : Munk sgaa rd . D av ie s , K .E . ( 1 986 ) e d . H uman G ene t ic D i sea se : a p rac t ica la pp roach . O xf o rd : I RL P re s s . d e V r ie s , R .R .P . , L a i , A . , F a t , R .F .M . , N i jenhu i s , L .E . &R ood , J . J . v an ( 1976 ) . HLA l ink ed g ene t ic c on t ro l o f h o s t r e spon se t o Mycobac te r ium L eprae . L ance t 2 : 1328 -30 . G ü r t le r , L .G . , J age r , V . , G rube r , W . , H i l lma r , I . , S chob loch , R . , Mü l le r , P .K . & Z iege lmeye r , G . ( 1 981) . P re senc e o f p ro te in s i n h uman b one s 2 00 , 1 200 , 1 500 y ea r so fa ge . H um .B io l .5 3 :137 -50 . H age lbe rg , E . , S yke s , B . , & H edge s , R . ( 1989 ) . A nc ien tb one DNA a mp l if ied . N a ture 3 42 :485 . H a r t , G .D . ( 1980 ) . A nc ien t d i sea se s o ft h e b l ood . I n B lood P ure a nd E loquen te d .M .W in t rob e ,p p . 3 3 -55 . N ew Y o rk : McG raw -H i l l . H edge s , R .E .M . a nd Wa l lace C a rm icha e l , J .A . ( 1978 ) . T he s u rv iva l o f b i ochem ica l i n fo rma t ion i n a r chaeo log ica lb one . J .A rchaeo l .S c i .5 : 377 -86 . J one s , J . S . ( 1 986 ) . Mumm i f ied h uman DNA c l oned . N a tu re 3 14 :576 . L owen s te in , J .M . ( 1980 ) . S p ec ie s -spec if ic p ro te in s i n f o s s i l s . Na turw i ss .6 7 :343 -46 . L owen s te in , J .M . ( 1981 ) . I mmuno log ica l r e ac t ion s f r om f o ss i l m a te r ia l . P h i los . T ran s . S oc . L ondon B 2 92 :143 -49 .

A cknow ledgemen ts Is hou ld l i ke t o t h ank D r . L . C a s te l le t t i a nd M r . A . Ma sp e ro o f t h e Mu seum a nd L abo ra to ry o fA r chaeob io logy o fC omo , I t a ly , f o rp e rm i s s ion a nd a s s i s tan ce i nu s ing t h e c ompu t e r f a c i l i t ie s . I a m i n deb ted a l so t oD r . K . G e l s tho rpe a nd D r . R . S oko l o f t h e R eg iona l B lood T r an s fu s ion C en t r e i nS he f f ie ld f o r t h e i r u n f a i l ing a s s i s tance d u r ing t h e p ro je c t o n t h e e x t ra c t ion o f HLA a n t igen s f r om a nc ien t h uman b one .

4 5

TABLE 5 . 1 L IST OF HLA ANT IGENS . ( AFTER B ENDER , 1 984 )

HLA -A A l A 2 A 3 A 9 AW23 AW24 A l0 A 25 A 26 AW24 A ll A 28 AW 19 A 29 AW30 AW31 AW32 AW33 AW34 AW36 AW43

HLA -DR DR W 1 DR W2 DR W3 DR W4 DR W5 DR W6 DR W7 DR W8 DR W9 DR W 10

HLA -B B 5 BW51 BW52 B 7 1 3 8 BW59 B 12 BW44 BW45 BW59 B 13 B 14 B 15 BW62 BW63 BW 16 BW38 BW39 B 17 BW57 BW58 B 18 BW2 1 BW49 BW50 BW22 BW55 BW56 B 27 BW35 B 37 B 40 BW60 BW6 1 BW4 1 BW42 BW46 BW47 BW48 BW53 BW54 BW4 BW6

HLA -DC DC ! DC3

HLA -C CW 1 CW2 CW3 CW4 CW5 CW6 CW7 CW8

HLA -BR B R3 B R4

HLA -D DW 1 DW2 DW3 DW4 DW5 DW6 DW7 DW8 DW9 DW 10 DW 11 DW 12

HLA -SB S B1 SB2 S B3 SB 4 SB

TABLE 5 . 2A

HLA -A , B , C ALLELE FREQUENCIES ( % ) .( AFTER BENDER , 1 984 )

C aucas ians E urope U SA

A l A 2 A 3 AW23

N egro es U SA A f r ica

1 5 .8 2 7 .0 1 2 .6

1 6 .1 2 8 .0 1 4 .1

8 . 1 1 6 .3 7 . 0

3 . 9 9 . 4 6 . 4

8 . 8 2 . 0

7 . 3 2 . 6

5 . 1 0 . 4

2 . 43 3 . 5

3 . 9 5 . 1 4 . 4 5 . 8 3 . 9 2 . 3 2 . 9 0 . 7 0 . 0 2 . 2

3 . 4 5 . 1 4 . / 3 . 6 2 . 9 4 . 5 3 . 7 1 . / 0 . 0 1 . 3

2 . 3 2 . 8 5 . 8 2 . 3 1 3 .0 2 . 8 1 . 9 5 . 1 0 . 0 1 6 .5

4 . 5 0 . 0 8 . 9 6 . 4 2 2 .1 4 . 2 1 . 5 1 . 0 0 . 0 1 .0

5 . 9 1 0 .4 9 . 2 1 6 .6 3 . 2 2 . 4 6 . 2 4 . 6 4 . 8 2 . 0

5 . 9 1 0 .5 1 0 .4 1 3 .8 2 . 6 5 . 1 3 . 1 5 . 6 5 . 9 2 . 5

4 . 9 1 / .6 5 . 5 1 4 .0 0 . 4 4 . 6 3 . 6 0 . 8 4 . 7 0 . 4

3 . 0 7 . 3 7 . 1 1 2 .7 1 . 5 3 . 6 2 . 0 0 . 0 3 . 0

3 . 5 5 . 7 2 . 2 3 . 6 9 . 9 1 . 1 8 . 1 0 . 0 0 . 0 3 . 6

1 . 4 4 . 9 3 . 8 / 3 8 . 6 1 . 7 9 . 2 0 . 0 0 . 0 2 . 8

0 . 4 1 .2 4 . 4 3 . 9 1 / .5 1 .) 3 . 9 0 . 0 0 . 0 1 .0

A 9 AW24 A 25 A 10 A 26 A ll A 28 A 29 AW30 AW31 AW32 AW33 AW43 B lank

O r ien ta l s J apan U SA

1 . 2 2 3 .3 0 . 7

2 . 5 4 3 .3 0 . 6

3 7 .2

2 3 .2

1 2 .7

0 . 6

6 . 7 0 . 0 0 . 2 0 . 5 8 . 7 0 . 5 2 . 0 0 . 0 4 . 2

0 . 0 2 . 8 0 . 6 1 . 1 1 9 .9 1 . 1 . 0 6 0 . 0 1 . 8

HLA -B B 5 B 7 B 8 B 12 B 13 B 14 B 18 B 27 B 15 BW38 BW16 BW39 B 17 BW21 BW22 BW35 B 37 B 40 BW41 BW42 B lank

2 0 .9 7 . 1 0 . 2 6 . 5 0 . 8 0 . 5 0 . 0 0 . 3 9 . 3 1 . 8

1 . 5

C auca s ians E urope U SA

1 6 .1 1 . 5 0 . 0 7 . 2 0 . 0 2 . 0 1 . 5 1 2 .3 1 7 .9

N egro es U SA A f r ica

1 4 .0 0 . 6 1 . 7 1 . 7

0 . 6 6 . / 1 3 .7 1 4 .5

4 . 7 0 . 6 1 . 5 6 . 5 9 . 4 0 . 8 2 1 .8 0 . 0 0 . 0 7 . 6

0 . 0 0 . 0 0 . 6 2 2 .1 0 . 0 1 6 .6 0 . 0 0 . 0 7 . 8

O r ien ta l s J apan USA

HLA -C CW 1 CW2 CW3 CW4 CW5 CW6 B lank

4 . 8 5 . 4 9 . 4 1 2 .6 8 . 4 1 2 .6 4 6 .7

3 . 7 6 . 0 1 .4 1 0 .2 5 . 2 1 .3 5 2 .1

1 . 9 9 . 2 8 . 8 1 2 .9 1 . 4 0 . 0 6 5 .8

4 7

0 . 0 1 .4 5 . 5 1 4 .2 1 . 0 1 7 .7 5 0 .2

1 .1 1 . 4 1 6 .3 4 . 3 1 . 2 2 . 1 5 3 .5

1 0 .1 4 . 6 1 6 .6 2 3 .4 1 . 1 0 . 0 4 4 .2

TABLE 5 . 2B

SO ME HLA -A AND B ANT IGEN FREQUENCES (% ) AMONG EUROPEANS , MIDDLE EASTERNS , I NDIANS , MONGOLOIDS .( AFTER DAUSSET AND COLO MBAN I , 1 973 ) .

E uropean

M idd le E as tern

I nd ian

Mongo lo id

HLA A l A '. ) A 3 A 9 A 10 A ll A 28 A 29 AW23 AW24 AW25 AW26 AW30

3 1 4 8 / 8 1 7 1 2 1 / 8 8 4 1 4 4 8 4

2 4 3 6 1 9 3 4 1 0 1 ' ) 1 ' ) / 1 / / 4 / 8 8

2 3 2 8 1 4 2 6 1 4 3 3 1 2 2 4 2 3 2 1 2 2

4 3 3 / 6 5 1 4 / 4 4 / 4 6 3 4 1 0 4

1 3 5 B 7 B 8 B 12 B 13 B 14 B 27 BW35

1 2 2 4 2 1 2 9 4 8 8 1 9

2 8 6 6 1 7 6 8 4 2 4

3 4 1 0 1 5 1 9 4 4 2 3

1 7 4 2 6 8 6 1 /

4 8

TABLE 5 . 3 SO ME S IGNIF ICANT HLA ANT IGEN AND D ISEASE A SSOCIATIONS

D i s ease

H LA

R p _e _

R heuma to id A r th r i t i s

B 27 DR4 BW54 DR4 DR4

C C 0 0 N

2 . 0 3 . 9 2 . 5 2 . 8 5 . 4

J uven i le R heuma to id A r th r i t i s

B 27 DR5

C C

3 . 9 3 . 3

A nky los ing S pondy l i t i s

B 27 B 27 B 27

C 0 N

6 9 .1 2 07 .9 5 4 .5

R e i t e r ' sD i sea se

B 27

C

3 7 .1

J uven i le D iabe t e s Me l l i tu s

B 8 B 15 DR3 DR4 BW54 DR3 DR4 B 8 B 15 DR3 DR4

C C C C 0 0 0 N N N N

2 . 6 2 . 2 3 . 4 3 . 7 5 . 7 4 . 9 2 . 6 2 . 5 2 . 2 3 . 2 6 . 7

G rave ' sD i sea se

B 8 DR3 B 35

C C 0

2 . 6 3 . 8 4 . 4

C e l ia cD i sea se

B 8 DR3 DR7

C 0 C

7 . 7 1 .7 7 . 7

P so r ia s i s V u lga r i s

B 13 B 17 B 37 CW6 DR7 B 13 B 17 B 37 CW6 DR7

C C C C C 0 0 0 0 0

4 . 1 5 . 3 4 . 0 7 . 5 3 . 3 3 . 3 2 . 0 8 . 5 8 . 5 7 . 7

I d iopa th ic H aemoch roma to s i s

A 3 B 7 B 14

C C C

6 . 8 3 . 0 2 . 8

S y s tem ic L upu sE ry thema to su s

B 8 DR3

C C

2 . 8 2 . 7

G oodpa s tu re ' s S ynd rome

DR2

C

1 3 .8

D e rma t i t is H e rpe t i fo rm i s

B 8 DR3

0 C

9 . 9 1 7 .3

Mu l t ip le S c le ro s i s

B 7 DR2

C C

1 . 9 2 . 8

Mya s then ia G rav i s

B 8

C

3 . 4

N a rco lep sy

DR2 DR2

0 C

3 58 .1 1 29 .8

C =C auca s ian

0 =O r ien ta l

R e la t ive r i sk

N =N eg ro

T he R e la t ive R i sk F ac to r i sa n i n d ica t ion o fh ow m uch m o r e o f ten t h e c a r r ie r o f a n a n t igon w i l l b e a f f ec ted b y t h e d i sea se , i . e . t h e r i sk o fd eve lop ing t h e d i sea se w i th in t h ec a r r ie ro f as p ec if ic a n t igen r e la t ive t ot h a t f o r i n d iv idua l s n o t c a r ry ing t h a t a n t igen ( Ber na l , 1 986 ) . Ar e la t ive r i sk g r ea t e r t h an 1i sp r e sen t w hen a n a n t igen i sm o re f r equen t i nt h e p a t ien t st h an i nt h ec on t ro l( Sve jgaa rd a nd R yde r , 1 977 ) . 4 9

L IFESTYLE , CULTURE CHANGE , POPULATION DYNA M ICS

HEALTH /D ISEASE

F igure 5 . 1

ENV IRONMENT

GENET ICS

b ac ter ia , c l ima te , p a laeoeconomy , d i e t , c u l tura l" hab i ts" , e t c .

i nna te p red ispo s i t ion t od i seas e

S cheme s how ing t h e i n t roduc t ion o fg ene t ic ( a long w i th e nv i ronmen ta l ) i n f o rma t ion i nt h er e con s truc t ion o ft h eh ea l th s t a tu s a nd t h e r efo re o ft h el i f e s ty le a nd d ynam ic so f as k e le ta lp opu la t ion .

H2

B2

m icrog lobu l in

COOH

CELL EXTER IOR

CELL MEMBRANE

CELL

I NTERIOR COOH

F igure 5 . 2

D raw ing o f ac l a s s IHLA a n t igen . C la s s IHLA a n t igen s h ave am o lecu la r w e igh to f4 5000 . E ach c ha in i si n se r ted i n to t h ec e l l m emb rane s ot h a t as ho r ts e gmen t i sw i th in t h e c e l l , o ne s egmen ta c ro s s t h e l i p id b i laye r o f t h e c e l l m emb rane , a nd t h e l a rge s t s e gmen t i se xpo sed t ot h e e x t er na l e nv i ronmen t a nd i s f o lded i n to t h ree d oma in s . C la s s Ia n t igen s m ak e u p c i rca 1 % o ft h e m emb rane p ro te in s . A s soc ia ted t o t h i sb i gge rs e gmen ti s as ma l lp r o te in , m ic rog lobu l in .( A f t e rA lbe r t se ta l . , 1 983 ) .

5 0

F igure 5 . 3

DOw2

DR3

B8

Cw7

A l

DO w3

DR 5

B35

Cw4

A ll

S chema t ic e xamp le o f t wo h ap lo type s i n a n i nd iv idua l . E ach h ou se s t h e A ,B ,C ,DQ ,DR l o c i , b u t t h e a l le le sm ay b eo fd i f f e ren t " t ype s " ,e . g . A la nd A l l . ( Af t e rS ch ieb l , 1 986 ) .

c h i ld 1

F igure 5 .4

c h ild 3

c h ild 2

A2 I

B44 1

A ll

B35

c h i ld 4

I nhe r i tance u n i t s i n af am i ly s eg reg t ion a na ly s i s . T he m a ter na l a nd p a t er na l h ap lo type s a r e s hown . T he se 4h ap lo type s c an r e comb ine t og i ve f ou rp o s s ib le c omb ina t ion s i n of sp r ing . E ach h ap lo type i si n he r i ted a s au n i t . ( A f te rB ende r , 1 984 ) .

5 1

A M ORE ACCURATE RECONSTRUCTION OF THE CULTURE AND DYNA M ICS OF ANCIENT POPULATIONS / 1 ‘ +o t h er e nv ironmen ta l a nd a r chaeo log ica ld a ta

SOUNDER BASE FOR THE RECONSTRUCT ION OF THE HEALTH S TATUS

GENET IC ( INNATE ) SUSCEPT IBILITY TO D ISEASE

EXTRACT ION OF GENETICALLY I NFOR MAT IVE MOLECULES FRO M SKELETAL MATER IAL

F igure 5 . 5

G ene ra l s cheme s how ing t h e u se o fg ene t ica l ly i n f o rma t ive m o lecu le s , e xt rac t ed f r om s ke le t a l r ema in s , i nt h er e con s t ruc t ion o ft h ed ynam ic s a nd c u l tu re o f ap opu la t ion .

5 2

6 .

DENTAL H ISTOLOGY A S AN I NDEX OF PAST CO M MUN ITY HEALTH

S imon H i l l son e xamp le ,l e ng th s o fl o ng b one s i nt h e l i mb s . I ti s a l so p o s s ib le t oo b ta in a r e a sonab le e s t ima te o f a ge -a t -dea th i n c h i ld r en a nd p o ten t ia l ly t h e m ea su r emen t sc ou ld b ep l o t ted a ga in s tt h e se a ge s . T he d i f f i cu l ty i s t h a t a l l t h e a ge e s t ima t ion m e thod s c ommon ly u s ed o n t h e r ema in s o f c h i ld ren a r e t h em se lve s b a sed o n p henomena r e la ted t ot h eg row th p ro ce s s . A ny r e l iab il i ty t h a t t h ey h ave a sa ge e s t ima to r s r e l ie s , i nf a c t , o n t h e a s sump t ion t h e r e i sn o d if erence i ng row th r a te b e tween a o nce l iv ing g roup o f p eop le r e p r e sen ted b y a r chaeo log ica l m a te r ia l a nd t h e m ode rn r e f e r ence c ommun i ty . I fg row th r a te i st o b e s t ud ied i na r cha eo logy , t h en s ome i n dependen t m ea su r eo fa ge i sr e qu i red .

C ommun i ty H ea l th T he c oncep to fh ea l th m igh tb es umma r i sed i n t e rms o f a c ommun i ty ' s d i e ta ry s t a tu s , t h e i n c idence o f i n f e c t iou s a nd o t he r d i sea se , o r s o c ia l c ond i t ion s w h i ch m ay i mp inge o n b o th m en ta l a nd p hy s ica l w e l l b e ing . I t i s v e ry d i f f i cu l t t od e f ine ag ene ra l i n dex o fh ea l th , e ven i n l i v ing h uman c ommun i t ie s . Mo s t m a jo r s t ud i e s u se t h e g r ow th r a te o f c h i ld r en i n t h e c ommun i ty a s a t l e a s t a g ene r a l i n d i ca to r . G row th r equ i re s as u rp lu si nt h eb ody ' sr e sou r ce s , o ve r a nd a bove w ha t i sr e qu i red f o r d ay to -day m a in t enance . G row th r a te i st h e re fo r e l i m i t ed b y d i e ta ry d ef i c ienc ie s ( wh ich f a i l t o s upp ly t h e r e qu i red s u rp lu s ) a nd d i sea se s ( wh i ch c au se t h e b ody t o u se u p i t s s u rp lu se s i n f i gh t ing t h em ) . E ven e mo t iona l s t r e s s m ay d ec r ea se g r ow th r a te ( Ha r r i son e ta l , 1 977 ) . W i th l i v ing c ommun i t ie s , e s t ima te s o fg row th r a t e a r e c a lcu la t ed i nv a r iou s w ay s a nd t h en c ompa r ed w i th g row th r a te i na r e f e r ence c ommun i ty w h ich i ss upp l ied w i th a n a bundan t d i e t a nd h a s g ood h ea l th c a re -u sua l ly i n t h e U SA o r E u rope . T h i s r e f e rence d a ta i s a c cumu la ted o ve r al o ng p e r iod b y s t udy ing c h i ld r en w hen t h ey v i s i t c l in ic s a t r e gu la r i n t e rva l s . I n d iv idua l c h i ld r en a r e f o l lowed t h rough t h e i r l i ve s , m ea su r ing s t a tu r e , h ead c i r cum f e rence , w e igh ta nd ah o s to fo t he r g ene ra l i n d ice s o f o ve ra l l b ody s i ze . L a rge n umbe r s o f c h i ld r en s upp ly t h e d a ta f o r c ha r t s w h i ch s umma r i se t h e v a r ia t ion i n s i ze a ch ieved a t p a r t i cu la ra ge si nt e rm so fp e r cen t i le s . C ha r t sl i ke t h e se a r e f r equen t ly i s sued t op a ren t s s ot h a tt h ey c an p l o tt h e i ro wn c h i ld ' sp rog r e s sa nd s e ei fh eo r s h e l a g s s e r iou s ly b eh ind . When t h e h ea l th o f a no the rc ommun i ty i st ob e a s se s sed ,f o re xamp le i n aT h i rd Wo r ld c oun t ry , i t i s u sua l ly n o t p o s s ib le t o f o l low i n d iv idua l c h i ld r en o ve r a p e r iod o f y ea r s . A m u ch s ho r te r t i me s ca le i s n eeded a nd s o g row th r a te i se s tab l i shed b y a n a l ter na t ive p ro cedu re . L a rge s amp le s o fc h i ld r en a r e m ea su red a nd t h e i r a ge s a r e d e t e rm ined b y a s k ing t h e i r p a ren t s , e xam in ing m ed i ca l r e co rd s e t c . A ge c an t h en b ep l o t ted a ga in s ts i ze i n aw ay t h a t m ay b e c ompa red w i th t h e p e r cen t i l e s o ft h e r e f e r en ce c ha r t s . T he re a r e c l ea r ly l a rge p o s s ib i l i t ie s f o r e r ro r i nt h is a pp roach , b u t i th a s b een u s ed w i th s u cce s sf o rm any y ea r s . I n a n i d ea l w o r ld , i tw ou ld b e p o s s ib le t o a pp ly s im i la r m e thod s t o a r chaeo logy , b u t a ny a t t emp t t o d o t h i s i n s tan t ly i n vo lve s ac i r cu la r a r gumen t . I ti sp o s s ib le t ot a ke m ea su r emen t so f t h e s k e le ton w h ich r e f lec t o ve ra l l b ody s i ze -f o r

L ayer ing i nT ee th I n dep enden t c l o ck s d o e x i s t i n h uman p hy s io logy . Many d i u rna l a nd o t he r r hy thms r e la t ing t o t h e g r ow th p ro ce s s h ave b een d i scove r ed ( Dean , 1 987 ) . O n ly t h e h a rd t i s sue so f t h e b ody s u rv ive o n mo s t s i te s a nd t h e b e s t e xamp le so fg r ow th r h y thm s a mong s tt h e se o ccu r i nt h e t i s sue so ft h e t e e th . T he e name l t h a tf o rm s t h e o u te r j a ck e t o f t h e t o o th c r own , a nd t h e d en t ine w h i ch m ake s u p t h e m a in b ody o f t h e t o o th a nd r oo t ,b o th b ea rr e co rd so fd a i ly r hy thm s a nd o f r h y thm s w i th l o nge r p e r iod ic i t ie s i n t h e f o rm o fm i c ro s cop i cl a ye red s t ruc tu re s .

a )D en t ine l a yers T h in s e c t ion s o f d en t ine , v i ewed u nd e r t h e p o la r i s ing m ic ro s cope , f r equen t ly s how a f i ne , r e gu la r p a t te rn o f l i ne s ( F igu r e 6 . 1 ) . T he se a r e u sualy k nown a s t h e i n c r emen ta l l i ne s o f v on E bne r , a f te r t h e a u tho r o f o ne o f t h e c l a s s i c p ape r s d e s c r ib ing t h em , b u t t h ey m ay a l so b e d e s c r ibed a s A nd re sen ' s l i nes ( Kawa sak i e t a l , 1 980 ) o r Mummery ' s l i ne s ( S chm id t a nd K e i l , 1 971 ) . E xp e r imen ta l s t ud ie s w i th p i g s ( Y i lma z e t a l , 1 977 ) , u s ing r e gu la ri n jec t ion s o fm a rke rd ye s a tt wo -we ek i n t e rva l s , h ave s ugge s ted t h a t t h e se l i ne s r e p r e sen t ad a i ly ( 2 4 h ou r ) l a ye r ing o f d en t ine a ppo s i t ion . T he l i ne s v a ry b o th i n s p a c ing a nd p rom inen ce a nd i ns ome i n d iv idua l s t h e r em ay a l so b el i ne so fm a rk ed d i s to r t ion t ot h e s t ruc tu re o ft h ed en t ine m a t r ix , s pa ced a ti n t e rva l s i n t h e l a ye r ing s equence . T he se a r e u sua l ly k nown a s t h e c on tour l i nes o f Owen , a f te r t h e g r ea t a na tom i s t S i r R icha rd Owen w ho f i r s t d e s c r ibed t h em , b u t t h e r e i ss ome c onf u s ion o ve r t h e e xac t m ean ing o ft h i s t e rm a nd o ne a u tho r a t l e a s t p r e f e r s t o c a l l t h em S chr ege r l i ne s

5 3

c r y s ta l s g r ow o u tr a d ia l ly f r om i n d iv idua lc e n t r e s t of o rm s p he r ica l b od ie s w h ich g r ow u n t i l t h ey m ee ta nd f u s ew i th o ne a no the r( F igu re 6 . 1 ) . T h i s p a t t er n o ccu r s i n d ep enden t ly t o t h e c o l lagen f i b r i l s a nd o t he rs t ruc tu re s o ft h e o r ig ina l d en t ine m a t r ix b u t ,i nf r e sh d en t ine ,o r gan ic s t ru c tu re sa r e s t i l l p r e se rved i n s ide t h e f u l ly m ine r a l i sed t i s sue . E ven i fa l o r gan ic m a t te r i sr emoved c hem i ca l ly f r om f r e sh d en t ine ( a n " a no rgan ic " p r epa ra t ion ) , t h e o ve ra l l a r rangemen t o f o r gan ic c omponen t s m ay s t i l lb ev i s ib le u nde rt h em i c ro s cope b e cau se m ine ra l i sa t ion h a s t os ome e x ten t f o l lowed t h e i r s t r uc tu r e , i na dd i t ion t of o l low ing t h e d om inan t c a l co sphe r i t ic m ine ra l isa t ion ( S chm id t a nd K e i l , 1 9 71 ; J one s a nd B oyde , 1 984 ) . F ea tu r e s o f t h e o r ig ina l p r eden t ine m a t r ix m ay b e v i s ib le d ue t o t h i s p henomenon e ven i n a r chaeo log ica l s p e c imen s w h i ch h ave l o ng s i nce l o s t m o s t o f t h e i rc o l lagen . T he i n c r emen ta l l i ne s o f d en t ine r e p r e sen t b o th t h e p r o ce s s o f m a t r ix d epo s i t ion a nd m ine ra l i sa t ion . T hey m ay b e s t ruc tu r e s w h ich w e r e o r ig ina l ly f o rmed i nt h e p r eden t ine o r gan ic m a t r ix , a nd t h en p a r t ia l ly e mpha s i sed b y t h e p a t te rn o f m ine ra l i sa t ion t h a t f o l lowed t h em . T hey m ay a l so r e p r e sen t d e f i c ienc ie s o f m ine ra l i sa t ion w h ich o c cu r r ed i n dependen t ly t o t h e p r eden t ine l a ye r ing . T h is s i tua t ion i sm o s t c l ea r ly a ppa r en ta sb and s o fi n t erg lobu la rs paces ( F igu r e6 . 1 ) , w he r e t h e c a lco sphe r i te s h ave f a i led t oc oa le s ce , l e av ing az one o fp oo r ly m ine ra l i sed p r ed en t ine b e twe en . T he se h ave b een r e la ted e xpe r imen ta l ly t og r ow th d i s tu rb ing f a c to r s , s u ch a sf e ve ri nr a bb i t s ( Be rmann e ta l , 1 9 39 )a nd r a t s ( Ga r r i son , 1 9 40 ) , a nd d e f ic ienc ie s o fv i tam in s A a nd D i nd og s ( Me l lanby , 1 929 ) . A s t h e f r on t o f m ine ra l i sa t ion l a g s s ome 1 5p . b eh ind t h e f r on t o f m a t r ix f o rma t ion ( M jo r , 1 984 ) a nd d o es n o t a lway s f o l low t h e s ame l i ne , t h e p o s i t ion o f a b and o f i n te rg lobu la r s p ace s d oe s n o t c o in c ide w i th t h a t o f t h e p r eden t ine l a ye r w h ich w a s f o rmed a tt h e s ame t i me . T he c on tou r l i ne s o f Owen ( a bove ) , a s o r ig ina l ly d e sc r ibed , a r e f e a tu r e s o f t h e o r ig ina l p r eden t ine m a t r ix ( B rad fo rd , 1 967 ) . S ome t e x t s ( S chm id ta nd K e i l , 1 9 71 ) i n s i s t i n s t ead t h a t t h ey a r e l i ne s o f i n te rg lobu la rs p a ce s ,o t he r se qua te t h em w i th l e s s m a rk ed l i ne s o f p oo r m ine ra l i sa t ion ( Bha ska r , 1 9 86 ) a nd s t i l l o t he r s ( t en C a te , 1 980 ) r e po r tt h a t t h e t e rm i sc ommon ly u s ed t om ean e xagge r a t ed l i ne s o f v on E bne r . T he se l a s t a r e i n a n e ven g r ea te rs t a t eo ft u rmo i l . C a r efu lw o rk b yS chm id t a nd K e i l ( 1 971 ) h a s s hown t h a t v on E bne r ' s l i ne s a r e p a r t o f t h e o r gan ic m a t r ix a nd a r e t h e r e fo re f o rmed d u r ing t h e f i r s t p ha se o f d en t ine p r oduc t ion . T he p r oo f f o r t h is i st h a t , w hen t h e m ine ra l c omponen t o f f r e sh d en t ine i sr emoved c h em icaly ( a " d em ine r a l i sed " p r epa ra t ion ) , t h e l i ne s a r e s t i l l c l ea r ly v i s ib le . T hey a r e , h oweve r , a l so v i s ib le ( r a the rm o re f a in t ly )w hen t h eo r gan ic c omponen t i s r emoved , s o t h e m ine ra l i sa t ion m u s t , t os ome e x t en t , f o l low t h e o r gan i sa t ion o f t h e l i ne s i nt h e p r eden t ine m a t r ix . Wha t c ause s

( B rad fo rd , 1 967 ) . T he p rom inence o fb o th w ha tI s ha l l c a l l v on E bne r ' s l i ne s a nd t h e c on tou r l i ne s o f Owen i su nde r s ome k i nd o fs y s t em ic c on t ro l : i no ne i n d iv idua l ,p a t te rn so fl i ne sc an b em a tched b e tween a l t e e th w h ich w e r eb e ing f o rmed a tt h e s ame t i me . I ti s t h e r e fo re p o s s ib le t o m a t ch c o r r e spond ing t e e th f r om t h e r i gh t a nd l e f t s i de s o ft h ej aw , a nd f r om u ppe r a nd l owe rj aw s . I ti s a l so p o s s ib le t o m a t ch t e e th o f d i f f e r en t c l a s s w ho se d en t ine f o rma t ion t i me h app end s t o o ve r lap ( F igu re 6 . 2 ) . I n t h i s w ay i tm ay b e p o s s ib le t om a tch , f o r e xamp le , ac an ine w i th a f i r s t m o la r . T he f i r s tp r a c t ica l a pp l ica t ion o ft h i s p henomenon w a s b y t h e S wed ish f o r en s ic d en t i s t G ö s ta G u s ta f son ( 1 947 ) w ho , i n ab l ind t e s t , w a s a b le t o m a t ch u p t h e t e e th o f t h r ee d i f f e r en t i n d iv idua l s b y t h e se m eans . T he p a t te rn o f d en t ine l a ye r ing i sh i gh ly i n d iv idua l a nd c l ea r ly h a s p o t en t ia l a s am ean s o f i d en t if ica t ion i n f o r en s ic s t ud ie s . G u s ta f son d i d n o t s u gge s t c oun t ing t h el i ne so fv on E bne r ,b u tt h i sw ou ld b e al o g ica l e x ten s ion o f h i s w o rk . Ma tch ing o f d i f f e r en t t e e th i nt h e s e quence c ou ld p rov ide a n e x tended c h rono logy o f d en t ine f o rma t ion , v e ry m uch i nt h es ame w ay t h a tt r ee r i ng s e quence sd o . S uch a c h rono logy c ou ld e ven b e g i ven a n a c cu ra te s t a r t ing p o in t , b ecause t h ep o in t o fb i r th i sc l ea r ly m a rk ed b y ap rom inen t l i ne ( c a l led t h e n eona ta l l i ne ) i nt h e d en t ine o ft e e th w h ich w e re f o rm ing a tt h et i me ( S chou r , 1 9 36 ) . T he n eona ta l l i ne c an b e s e en i na l d ec iduou s t e e th a nd i na l a rge p r opo r t ion o f p e rmanen t f i r s t m o la r s . B y c oun t ing l a ye r s f r om t h e n eona ta l l i ne , i tw ou ld t h eo r e t ica l ly b e p o s s ib le t o d e te rm ine p r ec i se ly w hen e a ch t o o th i n t h e d en t i t ion s t a r ted a nd f i n i shed . I ft e e th w e re s t i l l b e ing f o rmed a tt h e t i me o fd ea th , t h e l a s t l a ye r f o rmed w ou ld y i e ld a n e xcep t iona l ly p r ec i se e s t ima te o fa ge -a t -dea th b u t , m o r e i mpo r tan t ly f r om t h e p o in t o f v i ew o f g r ow th s t ud ie s ,t h ec h rono logy o ft o o th f o rma t ion w ou ld a c ta sa n i n dex o fg r ow th r a t e w h ich c ou ld b e c ompa r ed w i th r a d iog r aph ic s t ud ie s o f d en ta l g r ow th r a te i nl i v ing p opu la t ion s ( e .g . Moo r r ee s e ta l , 1 963 ;A nd e r son e ta l , 1 976 ) . T he re a r e , h oweve r , c on s ide r ab le d i f f icu l t ie s e ven a t at h eo r e t ica l l e ve l . D i f f e r en t a u tho r s m ean d i f f e ren t t h ing s b y t h e t e rms t h ey u s e t o d e sc r ibe t h e l i ne s . E s t ima te s o f s p a c ing v a ry g r ea t ly a nd t h e e v idence f o r t h e d a i ly r h y thm u nde r ly ing t h e s t ruc tu re s s e en i nt h e m ic ro s cop e i sn o ta lway sc l ea r . P a r to ft h e t r oub le i st h a tt h e f o rma t ion o f d en t ine i n vo lve s t wo s e pa r a te p ha se s ,a nd i tm ay b ed i f f icu l tt od e te rm ine w h ich i sr e spon s ib le f o r ag i ven s t ruc tu r e o rl i ne i nt h e f u l ly f o rmed t i s sue . T he f i r s t p ha se o ff o rma t ion i s t h e d epo s i t ion o f a n o r gan ic m a t r ix c a l l ed p red en t ine . T h i sc on s i s t sl a rge ly o ff e l ted m a t so f t h e f i b rou s p ro te in c o l lagen , t o ge the r w i th o t he r o r gan ic m a te r ia lk nown a sg r ound s ub s tance . T he s e cond p ha se i st h e m ine ra l i sa t ion o f t h e t i s sue . T os ome e x ten tt h et i ny c r y s ta l so fm ine ra lf o l low t h e c o l lagen f i b r i l s , b u t t h e d om inan t p a t te rn o f m ine ra l i sa t ion i s c a lcosphe r i t ic . T ha t i s , t h e

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t h e l a ye r ing i nt h e p r eden t ine i sn o t c l ea r . S ome a u tho r s ( Schm id ta nd K e i l , 1 971 )s ugge s tt h a ti ti s d ue t o r e gu la r a l te rna t io n s i n t h e d i r ec t ion o f c o l lagen f i b r i l s , w h i l s t o t he r s ( J one s a nd B oyd e , 1 984 ) s ugge s t t h a t i t m ay r e p r e sen t c yc l ica l v a r ia t ion i nt h ep ropo r t ion o fg r ound s ub s tance t o c o l lagen . I na dd i t ion , i ti sn o t e n t i re ly c l ea rw h ich l i ne s s hou ld b e i d en t i f ied a s " l ine s o f v on E bne r " . Mo s tt e x tbook s g i ve as im i la rr a nge o ff i gu r e s f o r t h es p a c ing o ft h el i ne s : a bou t5 p .( O sbo rn , 1 981 ) , 3 . 5µ t o6 µ d ep end ing o n t h ep o s i t ion i nt h et o o th ( t en C a te , 1 980 ) , 4 t o 8 µ ( Bha ska r , 1 986 ) . B rad f o rd ( 1 967 ) d e f ined t wo t y p e s o f r e gu la r i n c r emen ta l l i ne , o ne o n a ve rage 2 5µ a pa r t a nd t h e o t he r a bou t 3 4µ a pa r t . K awa sak i e t a l ( 1 980 ) s uppo r ted t h i s v i ew w hen t h ey f o und m o r e p rom inen t l i ne s a t2 0µ i n te rva l s i nt h e d en t ine u nde rt h e t o o th c r own , w i th f ou rf a in t e ra nd m o r e i n te rm i t ten t l i ne s i nb e twe en t h em s p a ced a t4 µ i n te rva l s . T hey s ugge s t ed t h a t t h e o r ig ina l p ub l ica t ion s o f v on E bne r a nd A nd r e sen w e r e b o th d e sc r ib ing t h e c oa r se r i n te rva l o fl i ne s o n ly . S o w ha t s hou ld b e c a l led " l ine s o f v on E bne r "? I n m y o wn s e c t ions , b o th o f f r e sh ly e x t ra c t ed t e e th a nd a r chaeo log ica l s p ec imen s , t h e m o s t p rom inen t r e gu la r p a t te rn s o f i n c r emen ta l l i ne s h ave a s pac ing o f 1 8 t o 2 8µ i n t h e d en t ine i mmed ia te ly u nde r t h e c en t re o ft h e t o o th c r own . C a re f u le xam ina t ion o ft h ed en t ine b e tween m a jo r l i ne s i n my s e c t ions d oe s r e vea l i n d ica t ion s o f l i ne s b e tween 3 . 5µ a nd 5 µ a pa r t . T he m a jo r l i n e sd o ,h oweve r ,b e come m o re c l o se ly s p a ced a s t h ey a r e f o l lowed d own t owa rd s t h e s i de s o f t h e c r own . T he s i tua t ion i sf u r the r c onf u sed b y a t temp t s t ol i nk t h es pa c ing o ft h el i n es t oe s t ima t e sf o rt h e r a t e o f f o rma t ion o f d en t ine . T he l i n e s t h a t Y i lmaz e ta l( 1 977 , s e ea bove )r e la t ed t o ad i u rna l r h y thm i n p i g s a ppea r t ob e a bou t 6 . 25µ a pa r t w hen s ca led f r om t h e i r p ub l i shed m i c ropho tog raph . O the re xper imen t s b a sed u pon t h e i n jec t ion o f m a rke r d y es i n to e xp e r imen ta l a n ima l s h ave p roduced e s t ima te s f o r d en t ine a ppo s i t ion r a te r a ng ing f r om 1 6µ p e r2 4 h ou r s i n r a t s , t o4 p . i nr h e su s m onkey s ( Symon s , 1 967 ) . S ome d r ugs g i ven t h e rapeu t i ca l ly i n m an a l so p roduce a m a rk e r l i ne i n t h e d en t ine a nd t h e s e h ave a l so y i e lded e s t ima t e s o fa r ound 4 µ p e r 2 4 h ou r s i nt h e d e c iduou s t e e th o f c h i ld r en ( S chou r a nd P on che r , 1 937 ) . K awa sak i e ta l( 1 980 ) u s ed ad i f f e r en t t e chn ique , c ompa r ing t h e d en t ine l a ye r s w i th i n c remen ta l s t ruc tu r e s i n t h e n e ighbou r ing e name l w h i ch a r e t h ough t t o r e p r e sen t d a i ly g r ow th ( s e e b e low ) . T hey a l so s ugge s t ed t h a t , u nde r t h e c en t ra l p a r t o f t h e c r own ,d en t ine g row sa t ar a t eo f4 µ p e rd ay . O n t h eb a s i so ft h i se v idence ,i ts e ems l i ke ly t h a t i ti s t h e f i ne r i n c remen ta l l i ne s o f t h e d en t ine w h ich r e p r e sen t a2 4h ou rg row th r h y thm . I n s p i te o f t h e c on s id e rab le p o ten t ia l o f d en t ine l a ye r ing , ag r ea t d ea lo fb a s ic h i s to log i ca l r e sea r ch i sn eed ed b efo r e t h i s t e chn ique i sl i ke ly

t ob e ap r a c t ica l p ropo s i t ion , e ven i nf r e sh , w e l l p r e se rved m a te r ia l . b ) E na me l l ayers D en ta l e name l i sa tp r e sen t a mo re r e a l i s t ic p r opo s i t ion b ecau se , b e ing a lmo s t w ho l ly m ine ra l , i ti su sua l ly m u ch b e t te r p r e se rved t h an d en t ine i n a r chaeo log ica l c on tex t s . E name l l a y e r ing i sa l so r a the r c l ea re r , mo r e c ons i s ten t ly v i s ib le a nd p r e sen t s f ewe r d i f f i cu l t ie s o f i n te rp r e ta t ion . U nde r t h e s c ann ing e l ec t ron m ic ro s cope , f r a c tu r ed s u r f a ce s w h i ch h ave b e en e t ched f o r af ew s e cond s i na c id s how t h e m a jo r f e a tu r e s o f e name l s t ruc tu r e w e l l . T he mo s t p r om inen t f e a tu r e s a r e b und le s o f t i ny c r y s ta l s , a bou t 5 µ a c ro s s a nd c a l led p r ism s . T he p r i sms a l t e rna te ly s we l l a nd c on s t r ic t a l ong t h e i r l e ng th , w i th ar e gu la ri n te rva lo fa r ound 4 µ ( F igu re 6 . 3 ) . T h i s f e a tu re c an a l so b e s e en i nt h in s e c t ion s o f e name l , e xam ined u nde r t h e l i gh t m ic ro s cope , a s a l t er n a te l i gh t a nd d a rk m a rk ing s a l ong t h e p r i sm s . T hey a r ek nown a sp r i sm c r o s ss t r ia t ions a nd t h e r e i sn ow l i t t le d oub t t h a t t h ey r ep r sen t a d a i ly r hy thm ( Dean , 1 987 ) . C oun t s o f c r o s s s t r ia t ion s b e tween p rom inen t m a rke r s i n t h e e name l h ave b e en m a t ched w i th s im i la rc oun t s o f t h e l i ne s o f v on E bne r b e twe en c o r r e spond ing m a rke r si nd en t ine ( Kawa sak i e ta l , 1 980 ) . E name l a l so s how s ar o ugh ly w eek ly r hy thm . T h i s i sm o s t c l ea r ly a nd c ons i s t en t ly s e en i nt h in s e c t ion s o f e name l u nde r t h e t h e o p t ica l m ic ro s cope a nd u sua l ly a ppea r s a s d a rk , f u zzy l i ne s c u t t ing d i agona l ly a c ro s s t h e b ounda r ie s o f t h e p r i sm s ( F igu r e 6 . 4 ) . T hese a r e n amed b rown s t r iae o f R e t z iu s , a f te r t h e S wed i sh a na tom i s t , A nde r s R e tz iu s , w ho p ub l i shed t h e c l a s s ic d e s c r ip t ion ( 1 837 ) . C ro s s s t r ia t ion s m ay b e c oun t ed b e tween n e ighbou r ing b rown s t r iae o f R e tz iu s a nd t h e c oun t v a r ie s b e tween s even a nd n i ne , w i th a n a ve r age i n mo s t s t ud ie s o f a bou t e i gh t ( Dean , 1 987 ) . T he v a r ia t ion i nc oun t s m ay w e l l b ed ue t oe xp e r imen ta l e r ro r , b ecau se b rown s t r ia e a r e r a r e ly c l ea r -cu t f e a tu r e s . T hey t e nd t o b ecome i n d i s t inc t u nde r h i ghe r p owe r m agn if i ca t ion s a nd i n t h inne r s e c t ion s , w he r ea s c r o s s s t r ia t ion s b e come l e s s d i s t inc t i n t h icke r s e c t ion sa nd a tl owe rm agn i f ica t ion s . I tt h e re fo re s e em s r e a sonab le t o p o s tu la te a r o ugh ly e i gh t c r o s s s t r ia t ion , a nd t h e r e fo r e e i gh td ay , r h y thm t o t h e f o rma t ion o ft h e l a ye r s w h i ch a r e m a rked o u t b yt h eb rown s t r iae o fR e tz iu s . T he b rown s t r iae v a ry m a rked ly i n f o rm . S ome a r e m o r e p rom inen t t h an o t he r s . I n s ome c a se s t h ey h ave as ha rp l i ne a tt h e i rc en t re w h i ch a ppea r s t ol i nk u p al i ne o fc r o s s s t r ia t ions w h i l s t i no t he r c a se s , t h ey m ay b e v e ry d i f fu se f e a tu r e s . T h i s v a r ia t ion s e ems t ob ea ga in u nde rs ome f o rm o fs y s tem ic c on t ro l , b e cau se g roup s o fp rom inen t b r own s t r iae m ay b e m a t ched b e twe en t e e th i n o ne i n d iv idua l , a nd t h e c oun t s o f c r o s s s t r ia t ion s b e tween t h em m ay b e m a t ched a l so . T h i s w a s d emon s t ra ted b y F u j i ta ( 1 939 ) a nd w a s

5

i n ve s t iga ted f u r the r b y A nna -G r e ta G u s ta f son ( 1 955 ) a s am ean s o f m a t ch ing t e e th f r om o ne i n d iv idua l f o r f o r en s ic p u rpo se s . A s i nd en t ine , t h ep e r iod o fe name lf o rma t ion ( F igu r e6 . 5 ) v a r ie s b e twe en t e e th a nd t h e d i f f e r en t p e r iod s m ay b e a r ranged i na no ve r lapp ing s e r ie s f r om b i r th u p t o a r ound 1 5 y ea r s o f a ge . S t a r t ing f r om t h e n eona ta l l i ne , w h ich i s o f ten v i s ib le i n t h e p e rmanen tf i r s t mo la r , B oyde ( 1 963a ; 1 963b ) w a s a b le t ou se c oun t so fc r o s ss t r ia t ion sa nd m a t ch ing o fb rown s t r iae t oa r r ive a te xcep t iona l ly p r e c i se a ge e s t ima te s f o r i n d iv idua l s w ho h ad d i ed w h i l s t t h e i rt e e th w e re s t i l lf o rm ing . B u l l ion ( 1 987 ) w a s a b le , i n an umbe r o fc a se s , t op roduce s e quence s o f 3 50 o r m o re b rown s t r ia e w h ich r e p r e sen t ed s e ven y ea r s o f t o o th g r ow th i nc h i ldhood . I ti s p o s s ib le t oe s t ima te c r own f o rma t ion t i m ing f r om t h e se s e quence s a nd , a s i nd en t ine , t h i s c ou ld b e u s ed a s av e ry p r ec i se i n dex o fg r ow th r a te w h i ch c ou ld b e c ompa r ed b e tween d i f f e r en t a nc ien t c ommun i t ie s a nd w i th m oder nr e f e r ence m a te r ia l . I t i s p o s s ib le t o c on s t r uc t s u ch s e quen ce s r o u t ine ly b ecau se o f t h e g ood p r e se rva t ion o f e name l a nd t h e c l a r i ty o f b r own s t r ia e i n m o s t s e c t ion s . S i nce c oun t s o f c r o s s s t r ia t ion s i n e name l m ay b e m a t ched w i th c oun t s o f t h e l i ne s o f v on E bne r i n d en t ine ( Kawa sak i e ta l , 1 980 , s e e a bove ) i tm ay b e p o s s ib le t oc omb ine e name l a nd d en t ine s equence s t o c ove r p a t che s w he r e e i the r t i s sue i sp oo r ly p r e se rved o r t h e l a ye r ing l a ck sc l a r i ty . I n s p i te o f t h e i r p o ten t ia l , s u ch l a ye r s e quence s a r e r a re ly a t temp ted e ven i n e name l . T he m a in r e a son f o r t h i s m u s t b e t h e t i me c on sum ing a nd d e s t r uc t ive s e c t ion ing w h ich i s n eeded . Af u l l s equence w ou ld p r obab ly r e qu i r e f i r s t , s e cond a nd t h i rd mo la r s , ap r emo la r , c an ine a nd i n c i so ra l l t ob e s e c t ioned . E a ch n eed s t ob e e ncap su la t ed i n to a r e s in b l o ck ( H i l l son , 1 986 ) b e cau se a nc ien t t e e th a r e u sua l ly b r i t t le a nd s h a t t e r w hen s e c t ion ing i sa t temp ted . T he b e s t i mp r egna t ion p ro cedu r e s t a ke s e ve ra l w eek s a nd s e c t ion ing , s eve ra l h ou r s . I na dd i t ion t ot h i s , t h e t e e th a r e d e s t royed l a rge ly , a l though h i gh q ua l i ty r e p l ica s c an b e m ade b e fo r e s e c t ion ing . F o r a l t h e se r e a son s , s t ud ie s o f e name l l a ye r ing a r e f o cu s s ing p r ima r i ly o n t h e s u r f ace o f t h e t o o th c r own . T he b r own s t r ia e o f R e tz iu s m a rk o u t t h e p a t t e rn o f l a ye r ing i nt h e c r own ( H i l l son , 1 986 ) . T he f i r s t e name l l a ye r s a r e l a id d own u nde r t h e c u sp so rh i ghe s tr i dge so ft h ec r own a s as e r ie so f d ome s , o ne o n t o p o f a no the r . T he se d ome s i n c r ea se i na r ea a nd o ve r lap o ne a no the rd own t h e s i de s o f t h e c r own . When t h e f u l l h e igh t o f t h e c u sp s o r r i dge s h a s b een a t ta ined , t h e p a t t e rn c hange s t os l eeve l ike l a ye r s , l a id d own o n t o po f t h ed ome sa nd a ga in o ve r lapp ing o ne a no the ri na s e quence d own t h e c r own s i de s . I nt h i s w ay , t h e f i r s tm on th s o fe name l f o rma t ion a r e h i dden f r om t h es u r f ace u nde rt h el a s td ome shap ed l a ye rt ob e f o rmed . T he r ea f te r ,e a ch l a ye rr i se sa ta na ng le t o t h es u r f ace a nd l e ave si t sm a rk t h e r e . Whe r ee a ch

b r own s t r ia m ee t st h e s u r f a ce , af u r row i sf o rmed , r u nn ing a r ound t h e c i r cum f e rence o f t h e c r own ( F igu r e s 4a nd 6 ) . T he e dge o f t h e l a ye r i n b e twe en b rown s t r ia e i sr e p re sen t ed a s ar i dge b e twe en t h e f u r row s . Aw ho le s e r i e so ft h ese c an b e s e en i nu nwo rn t e e th , e x t end ing d own t h e f u l l h e igh t o f t h e c r own a nd l o ok ing l i ke as e r ie s o f w ave s , o r o v e r lapp ing s t ra ta . T he r i dge s a nd f u r row sv a ry g r ea t ly i nf o rm a nd h ave av a r ie ty o f n ame s ( R i sne s , 1 984 ) , b u t a r e t o g e the r n orma l ly c a l led p er ikyma ta . T h i s w a s t h e n ame g i ven t o t h em b y G us t av P r e i swe rk i n 1 895 . E a ch b rown s t r ia i st h ough t t ob er e p re sen ted b y t h e f u r row o f ap e r ikyma a nd e a ch f u r row i st h ough t t oe x tend f u l ly a r ound t h e c i r cumf e r ence o f t h e c r own ( R i sne s , 1 985a ; 1 985b ) . T hey m ay t h e r e fo re b e c oun t ed i n t h e s ame w ay a s b rown s t r iae a nd , a s sum ing t h e s ame e i gh t d ay a ve rage g row th r h y thm , m ay b e u s ed t oe s t ima te t h e t i me t a ken f o rt h ec r own s u r f ace t of o rm . E s t ima t e so fc r own f o rma t ion t i me e s tab l i shed b y t h e se m e an s h av e b e en u sed t o a r r ive a t i mp roved e s t ima t e s f o r t h e a ge a td ea th o f f o s s i l h om in id s ( B romage a nd D ean , 1 985 ; D ean e ta l , 1 986 ) . F u r the r p o ten t ia l l i e s i nt h e p o s s ib i l i ty o f b u i ld ing u pe x t ended l a ye rs e quen ce s ,a sw i th t h e l i ne s o f v on E bne r a nd t h e b rown s t r iae o f R e tz iu s . T h e p e r ikyma ta v a ry i n p rom inen ce , r e gu la r i ty a nd s p a c ing . P a r t o f t h i s i sr e la ted t o t h e w ay i nw h ich t h e c r own s g row : t h e g roove s o f t h e p e r ikyma ta a r e f u r the r a pa r t n ea r t h e o cc lu sa l t i p o f t h e c r own t h an t h ey a r e n ea r t h e c e rv ix ( whe r e t h e c r own m e e t s t h e r o o t ) . T h i s r e gu la r p rog r e s s ion i ss e en i na l l t e e th , b u t t h e re a r e a l so d e ta i led v a r ia t ion s w h ich a r e n o t p a r t o f i t . S ome p e r ikyma ta m ay b e p a r t icu la r ly p r om inen t a nd s e ve ra l o f t h ese m ay f o rm a p a t t e rn w h ichcan b e m a tched b e tween d i f f e r en t t e e th f r om o ne i n d iv idua l . B and s o f i regu la r p e r ikyma t a m ay a l so b e m a t ched b e tween t e e th a nd t h i s s ugge s t st h a tb o th a r e r e la t ed t os y s t em ic f a c to r si nt h es ame w ay t h a tp rom inence o fb rown s t r iae a nd l i ne so fv on E bne ra r e . P e r ikyma ta a r e a l so r e la ted t om a c ro scop i ca l ly v i s ib le d e f e c t s o f t h e c r own s u r f a ce . T he se t a ke v a r iou s f o rm s ,b u t a r e m o s t c ommon ly m a rked g roove s o rb and s o f p i t t ing w h i ch r u n r o und t h e c i r cum f e r ence o ft h e c r own a nd a r e l a rge e nough t o b e s e en b y t h e n aked e ye . S uch d e f e c t sh ave l o ng b e en o b se rved b y d en ta l s u rgeon s a nd o ne o f t h e k ey e a r ly s t ud ie s w a s b y R obe r t B unon i n 1 746 . B unon c a l led t h e d e f e c t s e r o s ion s a nd d emons t ra t ed t h em i nt h e u ne r up ted t e e th o f c h i ld r en w ho h ad s u f f e red f r om s u ch d i sea se s a s r i cke t s , s cu rvy , m ea s le s a nd s ma l lpox . D e f e c t s c ou ld h ave o c cu r r ed i n t h e se t e e th o n ly b y i n t e r r up t ions t o t h e g r ow th p r oce s s . T he l i nk w i th d e f ic ienc ie s i n v i tam in s A a nd D w a s e s tab l i shed b y L ady Me l lanby i n h e r e xp e r imen t s w i th b eag le d og s ( 1 929 ) a nd a v a r ie ty o f c l in ica l s t ud ie s i n m an h ave s i nce c on f i rmed ac onnec t ion w i th v a r iou s n u t r i t iona l d e f ic ien c ie s , i n f ec t iou s d i sea se s a nd o t he r f a c to r s ( P indbo rg , 1 982 ) . T he d e f e c t s a r e

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t h e i r p o t en t ia l s t i l l r e qu i re s ag r ea t d ea l o f h a rd w o rk o nb a s ic h i s to logy a nd p hy s io logy .

n owaday s r e f e r red t o a s e nam e l h ypop la s ia , m ean ing u nde r -deve lopmen t o f t h e e name l . I n s ome c a se s t h ey c an b e m a t ched w i th i n t e rg lobu la rs pace si nd en t ine ( a bove ) ,b u tt h i si s b y n o m ean s a lway s s o . E ven t h ough t h e i r g ene ra l r e la t ion sh ip t og r ow th d i s rup t ing f a c to r s s u ch a s f e ve r a nd d i e ta ry d e f ic iency s e em s c l ea r , i ti sd i f f icu l tt ol i nk p a r t icu la rd e f ec t st op a r t icu la r d i s tu rban ce s . S a rna t a nd S chou r ( 1 941 ; 1 942 ) m a t ched o n ly 5 0% o f d e f ec t s w i th e p i sode s o f f e ve ri nt h em ed ica lr e co rd so f6 0c h i ld r en . I na n u npub l i shed s t udy o f 2 8 y oung m en , Im a t ched 7 0% o f d e f ec t s w i th e p i sode s r e co rded i n t h e i r m ed ica l r e co rd s o n ly w hen t h e mo s t m ino r d e f e c t sw e re i n c luded . I n s p i te o f t h e d i f f i cu l t ie s i n e xp la in ing i n d iv idua l d e f ec t s i n l i v ing p eop le , t h ey a r e c l ea r ly o f s y s tem ic o r ig in a nd c an b e m a t ched , a l ong w i th i regu la r i t ie s o f p e r ikyma ta , b e tween t e e th i n o ne i n d iv idua l . O n c l o se r s t udy , t h e g r oove l ike d e f ec t s a r e t h emse lve s s e en t o b e f o rmed b y v a r ia t ion si nt h es pa c ing o fp e r ikyma ta ( F igu re 6 . 6 ; H i l lson a nd J one s , 1 989 ) . T he p e r ikyma ta /hypop la s ia s e quen ce s o f d i f f e r en t t e e th m ay t h e re fo re b e m a t ched u p t op rodu ce a n e x t ended s equence o ft h e k i nd s ugge s ted f o rl i ne s o f v on E bne r a nd b r own s t r iae o f R e tz iu s . T h i s w a s s ugge s ted b y m y se l f ( H i l l son , 1 979 ; 1 986 ) a nd h a s s i nce b een t e s ted b y o ne o fm y r e sea r ch s t uden t s ( Bu l l ion , 1 987 ) , w ho w a s a b le t o c on s t ruc t c ompo s i te s e quence s o f u p t o 2 00 p e r ikyma ta , r e p r e sen t ing a r ound f o u r a nd a h a l f y ea r s o fg row th . T he se s e quence s m ay b e l i nk ed a pp rox ima te ly t oc h rono log ica la ge i ft h ea ve rage a ge s f o r t h e s t a r t a nd c omp le t ion o f t h e h i dden d ome l ike l a ye r s a r e k nown f o r s ome o ft h e t e e th i n t h e s equence . B u l l ion ( 1 987 ) h a s m ade a p i onee r ing a t temp t a t e s tab l i sh ing t h e se f o r a r a nge o f t e e th , b y c oun t ing b r own s t r ia e o f R e t z ius i n t o o th s e c t ion s . P e r ikyma ta c an b e d i f f i cu l t t o s t udy a nd a r e q u i te r a p id ly l o s t b y a b ra s ion o f t h e c r own s i n a du l t t e e th . T he p o ten t ia l , h oweve r , i s g r ea t b e cau se b o th p e r ikyma ta a nd h ypop la s ia c an b e e xam ined n on d e s t r uc t ive ly i nc a s t so ft h ec r own s u r f a ce . T he se c a n b e m ade f r om t h e t e e th o f l i v ing p eop le a s w e l la sa nc ien tb u r ia l s . T h ey m ay y i e ld t h e s ame k i nd o f i n fo rma t ion o n d en ta l g r ow th r a t e a nd g r ow th d i s tu rbance s a s t h e b r own s t r ia e o f R e tz ius a nd t h e i n c r emen ta l l i ne s o f d en t ine , s o t h a t t h ey o f f e r t h e b e s t h ope o f ag r ow th i n dex t h a tc ou ld b e d e t e rm ined r o u t ine ly a nd c ompa r ed w i th l i v ing c ommun i t ie so fp eop le .

R e f erence s A nde r son , D .L . , T homp son , G .W . & P opov i tch , F . ( 1976 ) . A ge o fa t ta inmen to fm ine ra l i sa t ion s t age so f t h ep e rmanen td en t i t ion . J .F o r .S c i .2 1 :191 -200 . B e rmann , M . , E dwa rd s , L . & K i t ch in , P . ( 1939 ) . E f f ec t s o f a r t if ic ia l ly i nduced h ype rpy rex ia o n t o o th s t ruc tu re i n t h e r abb i t . P r o c . S o c . E xp . B io l . M ed . 4 1 :113 -15 . B ha ska r , S .N . ( 1986 ) . e d . O rb an ' s O r a l H i s to logy a nd E mb r yo lo gy . S t .L ou i s : C .V . Mo sby . B oyde , A . ( 1963a ) . E s t ima t ion o f a ge a t d ea th o f y oung h uman s k e le ta lr ema in s f r om i n c remen ta l l i ne s i n t h e d en ta l e name l . 3 rd I n t e r na t iona l M e e t ing i n F o r en s i c I mmuno lo gy , M ed ic in e , P a t ho logy & T ox i co lo gy ( L ond on A p r i l1 6 t h -24 th 1 963 )P l ena ry S e s s ion I A . B oyde , A . ( 1963b ) . U npub l i shed d a ta q uo ted b y M i le s , A .E .W . I n D en t a l A n t hropo log y e d . D .R . B ro thwe l l . p p . 1 9 1-2 10 . L ondon : P e rgamon P re s s . B rad fo rd , E .W . ( 1967 ) . M ic roana tomy a nd h i s tochem i s t ry o f d en t ine . I n S t ru c tu r a l a nd C hem ic a l O r gan i z a t ion o fT e e t h . e d . A .E .W . M i le s . p p . 3 -32 . O r lando : A cadem ic P re s s . B romage , T .G . a nd D ean , M .C . ( 1985 ) . R e -eva lua t ion o f t h e a ge a t d ea th o f i mma tu re f o s s i l h om in id s . N a tu r e3 /7 :525 -27 . B ro thwe l l , D .R . ( 1963 ) . e d . D en ta l A n thropo lo gy . L ondon : P e rgamon P r e s s . B u l l ion , S .K . ( 1 987 ) . I n c r emen t a l S t ruc tu r e s o fE name l a nd t h e i r A pp l i ca t ion s t o A r chaeo logy . U npub l i shed P h .D .t h e s i s ,U n ive r s i ty o fL anca s te r . D ean , M .C . ( 1987 ) . G row th l a ye r s a nd i n c remen ta l m a rk ing s i nh a rd t i s sue s : ar e v iew o f t h e l i t e ra tu re a nd s ome p r e l im ina ry o b se r va t ion s a bou t e name l s t r uc tu re i nP a r an t h ropu s b o i s e i . J .H um . E vo l . 1 6 :157 7 2 . D ean , M .C . , S t r inge r ,C .B . &B romage ,T .G . ( 1986 ) . A n ew a ge a t d ea th f o r t h e N eande r tha l c h i ld f r om D ev i l ' s. T owe r , G ib ra l ta r a nd t h e i mp l ica t ion s f o r s t ud ie s o f g ene ra l g row th a nd d eve lopmen t i n N eand e r tha ls . A m .J .P hy s .A n t h r opo l . 7 0 :30 1 -09 . F u j i ta , T . ( 1 939 ) . N eue f e s t s te l lungen ü be r d i e R e tz iu s ' shen p a ra l le l s t r e i f en d e s Z ahn schme lz e s . A na t .A n z .8 7 :350 -55 . G a r r i son , L . ( 1940 ) . T he e f f ec t o f f e ve r o n t h e d eve lopmen t o f t h e r a t i n c iso r . J .D en t . R e s . / 9 :215 2 5 . G u s ta f son , A -G . ( 1 955 ) . T he s im i la r i ty b e tween c on t ra la te ra lp a i r so ft e e th . O don to l .T i d sk r .6 3 :245 -48 . G u s ta f son , G . ( 1 947 ) . M ic ro scop ic e xam ina t ion o f t e e th a s a m ean s o f i d en t i f ica t ion i n f o ren s ic m ed ic ine . J .A m .D en t .A s soc .3 5 :720 -24 . G u s ta f son , G . a nd K och , G . ( 1974 ) . A ge e s t ima t ion u p t o 1 6 y ea r s o f a ge b a sed o n d en ta l d eve lopmen t . O d on t o l .R e v y2 5 :297 -306 . H a r r i son , G .A . , We ine r , J . S . , T anne r , J .M . &B a rn ico t , N .A . ( 1977 ) . H uman B i o logy 2 nd e d i t ion . O xf o rd : O x fo rd U n ive r s i ty P re s s . Wi l son , S .W . ( 1979 ) . H uman B io lo g i ca l V ar ia t ion i nt h e N i l e V a l l e y , i n R e l a t ion t o E nv i ronmen ta l F ac to r s . U npub l i shed P h .D .t h e s i s , Un ive r s i ty o fL ondon . H i l l son , S .W . ( 1986 ) . T e e t h . C amb r idge : C amb r idge U n ive r s i ty P re s s . H in son , S .W . a nd J one s , B .K . ( 1989 ) . I n st rumen t s f o r m ea su r ing s u r f ace p ro f i le s : a n a pp l ica t ion i n t h e

C onc lus ion T he l a ye r ed s t ruc tu r e o f e name l a nd d en t ine o f f e r s w ha t i sp robab ly t h e o n ly p o s s ib i l i ty i n a r chaeo logy f o r s t udy ing g r ow th r a te i nr e la t ion t oa n i ndependen t m ea su r e o f t i me . T he se t wo t i s sue s a r e l i ke ly t op rove av a luab le r e sou r ce i n b o th a r cha eo logy a nd a n th ropo logy , b u t r e a l i s ing

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s t udy o f a nc ien t h uman t o o th c rown s u r f ace s . J . A r chaeo l .S c i ./ 6 :95 -105 . J one s , S . J . a nd B oyde , A . ( 1984 ) . U l t ra s t ruc tu re o f d en t in a nd d en t inogene s i s . D en t in a nd D en t inogen e s i s . e d . A . L inde . p p . 8 1 -134 . B oca R a ton : CRC P r e s s I n c . K awa sak i , K . , T anaka , S .& I s h ikawa , T . ( 1980 ) . O n t h e d a i ly i n c remen t a l l i ne s i n h uman d en t ine . A r ch . O r a lB io l .2 4 :939 -43 . L ind e , A . ( 1984 ) . e d . D en t in a nd D en t ino gen e s i s . B oca R a ton : CRC P re s sI n c . Me l lanby , M . ( 1929 ) . D ie ta nd t e e th : a n e xp e r imen ta l s t udy . P a r t I . D en ta l s t ruc tu re i n d og s . M ed ic a l R e s ea r ch C ounc i l , S p e c ia l R epo r t S e r i e s N umb e r 1 40 . L ondon : HMSO . M i le s , A .E .W . ( 1963 ) . D en t i t ion i nt h e a s se s smen t o f i n d iv idua l a ge i n s ke le ta l m a t e r ia l . I n D en t a l A n t h ropo lo gy e d . D .R . B ro thwe l l . p p . 1 91-2 10 . L ondon : P e rgamon P r e s s . M i le s , A .E .W . ( 1967 ) . e d . S t ru c tu r a l a nd C hem ica l O r gan i z a t ion o fT e e t h . O r lando : A cad em ic P re s s . M jo r , I .A . ( 1984 ) . T he mo rpho logy o f d en t in a nd d en t inogene s i s . I nD en t in a nd D en t inogen e s i s . V o l . 1 . e d . A .L ine . p p . 1 18 . B oca R a ton : CRC P re s sI n c . Moo r r ee s , C .F .A . , F ann ing , E .A . &H un t , E .E . ( 1963 ) . A ge v a r ia t ion o f f o rma t ion s t age s f o r t e n p e rmanen t t e e th . J .D en t .R e s .4 2 :1490 -1502 . O sbo rn , J .W . ( 1981) . e d . D en ta l A na tomy a nd E mb r yo logy : A C ompan io n t o D en t a l S t ud ie s 1 ( 2 ) . O x fo rd : B la ckwe l lS c ien t if ic P ub l ica t ion s . P indbo rg , J . J . ( 1982 ) . A e t io logy o f d eve lopmen ta l e name l d ef ec t s n o t r e la ted t o f l uo ro s i s . I n t . D en t . J . 3 2 :123 -34 . R e tz iu s , A . ( 1 837 ) . B eme rkungen ü b e r d en i n ne ren B au d e rZ ä thne . A r ch i v .A na t .P hy s io l .4 86 -566 .

R i sne s , S .( 1984 ) . R a t iona le f o rc on s i s t ency i nt h e u se o f e name l s u r f ace t e rms : p e r ikyma ta a nd i mb r ica t ion s . S c and .J .D en t .R e s .9 2 :1-5 . R i sne s , S . ( 1985a ) . A s cann ing e l ec t ron m ic ro scop e s t udy o ft h e t h r ee -d imen s iona le x ten to fR e tz iu s l i ne s i nh uman d en ta l e name l . S cand . J .D en t . R e s . 9 3 :145 5 2 . R i sne s , S . ( 1985b ) . C i rcumf e ren t ia l c on t inu i ty o f p e r i l cyma ta i nh uman d en ta l e name l i n ve s t iga ted b y s c ann ing e l e c t ron m ic ro scopy . S cand . J .D en t . R e s . 9 3 :185 -91 . S a rna t , B .G . a nd S chou r , I .( 1941) . E name l h ypop la s ia ( c h rono log ic e name l a p la s ia ) i n r e la t ion t os y s tem ic d i sea se : ac h rono log ic , mo rpho log ic a nd e t io log ic c l a s s i f ica t ion . J .A m .D en t .A s so c .2 8 :1989 -2000 . S a rna t , B .G . a nd S chou r , I .( 1942 ) . E name l h ypop la s ia ( c h rono log ic e name l a p la s ia ) i nr e la t ion t os y s tem ic d i sea se : ac h rono log ic , mo rpho log ic a nd e t io log ic c l a s s i f ica t ion . J .A m .D en t .A s so c .2 9 :67 -75 . S chm id t , W . J . a nd K e i l , A . ( 1971) . P o l ar i za t ion M i c ro s cop y o f D en t a l T i s sue s . Ox fo rd : P e rgamon P r e s s . S chou r , I .( 1936 ) . N eona ta l l i ne i n e name l a nd d en t in o fh uman d e c iduou s t e e th a nd f i r s t p e rmanen t mo la r . J .A m .D en t .A s so c .2 3 :1946 -1955 . S chou r , I .a nd P onche r , H .G . ( 1937 ) . T he r a te o f a ppo s i t ion o fh uman e name l a nd d en t in a sm ea su red b y t h e e f f ec t s o f a cu te f l uo ros is . Am . J .D i s . C h i ld . 5 4 :757 -76 . S ymon s , N .B .B . ( 1 967 ) . D en t inogene s i s . I nS t r u c tu ra l a nd C hem i ca lO r gan i z a t ion o fT e e t h . e d . A .E .W . M i le s . p p .2 85 -324 . L ondon : A cadem ic P re ss . t e n C a t e , A .R . ( 1980 ) . e d . O r a lH i s to lo gy . S t . L ou i s : C .V . Mo sby . Y i lmaz , S . , N ewman , H .N . & P oo le , D .F .G . ( 1977 ) . D iu rna l p e r iod ic i ty o fv on E bne r g row th l i ne s i np ig d en t ine . A r c h .O r a lB io l .2 :51 1 -1 3 .

5 8

F igure 6 . 1

I n c remen t a l s t ruc tu re s i nt h e d en t ine o f ah uman c an ine f r om t h e s i t e o fK e rma , N ub ia ( c . 1 720 -1550 B .C . ) . Ma te r ia l f r om t h e D uckwo r th L abo ra to ry , U n ive r s i ty o fC amb r idge . V on E bne r ' s l i ne s a r e t h e f uzzy d a rk b and s r unn ing h o r izon taly a c ro s s t h e f i e ld o f v i ew ( one i s ma rked E -E -E ) . T he c a lco sphe r i t e s c an b e s e en a sd a rk , c l oud l ike m a s se s . T owa rd s t h e b o t tom o ft h e f i e ld o fv i ew , s ome a r eo u t l ined b yn a r row p a tche so fi n t e rg lobu la rd en t ine ( Ma rked I ) . T he s ha rp , s i nuou s l i ne sr unn ing u p a nd d own t h e p ho tog raph a r e t h e d en t ina l t u bu le s , n o t a n i n c remen ta l s t ruc tu re , b u t ama jo r f e a tu re o f d en t ine h i s to logy . P o l i shed t h in s e c t ion v i ewed b yp o la r i s ing m ic ro scopy . F ie ld w id th c .5 0µ .

5 9

4

8

1 2

1 6

F igure 6 . 2

R ange s o fd en t ine f o rma t ion t i me s f o rv a r iou s h uman p e rmanen tt e e th ( d a ta f r om G u s ta f son a nd K och , 1 974 ) . C ode s f o r t e e th : m l =f i r s tm o la r ; i= i n c i so r s , c= c an ine , p= p r emo la r s , m2 = s e cond mo la r , m 3 =t h i rd m o la r . S ca le i ny ea r sa f te rb i r th .

6 0

F igure 6 . 3

P r i sms i nt h e e name l o f ah uman f i r s t m o la r f r om t h e C oppe rga te s i t e , Y o rk . T he p r i sms r un u p a nd d own d i agona l ly a nd s how t h e a l te rna t e s we l l ing s a nd c on s t r ic t ion s ( p rom inen t i nt h e a r ea m a rked C ) a l ong t h e i r l e ng th w h ich a r e t h ough t t oc o r r e spond t oc r o s s s t r ia t ion s . F ra c tu red , e t ched s p ec imen , e xam ined b ys c ann ing e l ec t ron m ic ro scopy , u s ing s e conda ry e l ec t ron m ode . F ie ld w id th c .6 0µ .

6 1

F igure 6 . 4

B rown s t r iae o f R e tz iu s i nt h e e name l o f ah uman c an ine f r om K e rma , N ub ia ( c . 1 750 -1550 B .C . ) . Ma te r ia l f r om t h e D uckwo r th L abo ra to ry , U n ive r s i ty o fC amb r idge . T he s t r iae ( one i sm a rked R -R -R ) a r e r unn ing a lmo s t v e r t ica l ly u p t h e p ho tog raph , t o m ee t t h e s u r f ace o f t h e c r own w he r e s l igh t i n den ta t ion s m a rk ing t h e f u r row s o ft h e p e r ikyma ta ( ma rked P ) c an b e s e en . T he p r i sms c an b e s e en c u t t ing a c ro s s t h e l i ne o f t h e b r own s t r iae d i agona l ly . P o l i shed t h in s e c t ion e xam ined b y p o la r i s ing m ic ro scopy . F i e ld w id th c .2 50µ .

6 2

m3 1 2

1 6 F igur e6 . 5

R ange so fe name l f o rma t ion t i me s f o rv a r iou s h um an p e rmanen tt e e th ( d a ta f r om G u s ta f son a nd K och , 1 974 ) . C od e sf o rt e e th a si nF igu re 6 . 2 . S ca l ei ny ea r sa f t e rb i r th .

6 3

F igure 6 . 6

P e r ikyma ta o n t h e c r own s u r f ac e o f ah uman c an ine f r om t h e L ive rpoo l S t ree t /B road S t ree t s i te , L ondon . Ma te r ia l f r om t h e E nv i ronmen ta l A rchaeo logy U n i t , Mu seum o fL ondon . am a rked g roove l i ke d ef ec t o f e name l h ypop la s ia ( ma rked H -H -H ) c an b e s e en , a nd ap i t ted d e fec t i sp a ra l le l t oi t , t owa rd st h et o po ft h ef i e ld o fv i ew . S i l icone r ubb e ri mp re s s ion t a ken f r om t h es u r face , s pu t te rc oa ted i n g o ld a nd e xam ined b y s c ann ing e l ec t ron m ic ro scopy , u s ing s e conda ry e l ec t ron m ode a nd i n ve r ted v i deo . T h i si s at i me sav ing t e chn ique , w h ich g i ve s t h ei l u s ion o f ap o s i t ive i mage t ot h en ega t ive i mp re s s ion , a l though t h e s hadow s a r e n o t q u i te a st h ey w ou ld b e i ft h e c r own s u r f ace i t se l f w e re b e ing e xam ined . F i e ld w id th c .

6 4

PART 2

C a se S tud ie s

7 .

PATTERNS OF L IFE AND DEATH : THE PALAEODE MOGRAPHY OF MEDIEVAL YORK .

A nne L .G rauer A rme lago s ( 1 976 ) , P a lkov i ch ( 1 980 ) a nd K ennedy ( 1 984 ) t o m ak e s i gn if i can t i n f e rence s a bou t p a t ter n s o f n u t r i t iona l d e f ic ienc ie s a nd d i sea se a c co rd ing t oa ge ,s e x ,s t a tu s ,g eog raph ica l l o ca t ion a nd s ub s i s tence p a t t er n . B u t l i f e t a b le a na ly s i s h a s i t s p rob lems . T he d eba t e o ve r t h e a pp l icab i l i ty a nd b i a se s i n he ren t i n t h e u s e o f s k e le ta l m a te r ia l h a sr e cen t ly b ecome h ea t ed ( s ee A nge l , 1 969 ; B o cque t -Appe l a nd Ma s se t , 1 982 , 1 985 ;a nd H owe l l , 1 982 f o ra r gumen t sa ga in s tt h e u s e o fp a laeodemog raphy a sa c cu ra te r ef l ec t ion s o f a r chaeo log ica l p opu la t ions , a nd V an G e rven a nd A rme lago s , 1 983 ; B u ik s t ra a nd K on ig sbe rg , 1 985 ; a nd G re en e ta / , 1 986 f o r a r gumen t s i n f a vou r o f c a re f u l a pp l ica t ion o f p a laeodemog raph i c d a ta t o a r chaeo log ica l m a te r ia l ) . T he t h ru s t o f t h e c r i t i c i sm a ga in s t l i f e t a b le a na ly s i sh a sb e en a imed a tt h eu s eo fa b so lu te a ge a td ea th c a lcu la t ion s f r om s k e le ta l m a te r ia l , t h e i n ab i l i ty o ft h er e su l t st or e f le c tm o r ta l i ty p a t ter n s o f i n tended p opu la t ion s ( t he m o r ta l i ty c u rve m im ic s t h er e f e r ence p opu la t ion e mp loyed t oa ge t h e s k e le ta l m a t er ia l ) , t h e n e ce s s i ty t o a s sume s t ab le p opu la t ion d ynam ic s ,a nd t h el a ck o fl i v ing p opu la t ion sw i th s im i la rd emog r aph i c s t ruc tu r ea s t h o se c r ea ted f r om s k e le ta lp opu la t ion s . I n r e fu ta t ion , t h e se c r i t ic i sms h ave b e en c ha l lenged b y t h e u s e o f m u l t ip le t e chn ique s o f a ge ing a nd s e x ing s k e le ta l m a te r ia l ( Love joy e t a l , 1 985 ) , w h ich r e ly o n t h e c oncep t o f d eve lopmen ta l a ge s r a the r t h an a b so lu te a ge a t d ea th . A l so i nr e f u ta t ion o ft h ec r i t ic i sm ,G r eene e t a l ( 1 986 ) , u s ing We i s s ' ( 1 973 ) mode l l i f e t a b le s , s how t h a t t h e i r s k e le ta l s amp le s f r om K u lubna r t i " a re m o r et h an c op ie so fa ny s o -ca l led r e f e r en ce p opu la t ion . T he p a t te rn s , i n s tead , a r e a r ef l ec t ion o f o ne s e t , a mong m any , o f r e a l a nd b i o log ica l a nd c u l tu ra l p ro ce s se s a f f ec t ing l i f e a nd d ea th " ( G reene e ta l , 1 986 :199 ) . T he n e ce s s i ty t o a s sume s t ab le p opu la t ion d ynam ic sh a sa l so b een a dd r es sed b y Moo re e ta l ( 1 975 ) a nd J ohan s son a nd H o row i tz ( 1 986 ) . A s sum ing f i r s t t h a t ap opu la t ion i ss t ab le a l low s t h e r e sea r che r t oe xp lo r e t h e e f f ec t s o f p o s s ib le i n s tab i l i ty i n t h e p opu la t ion . F o r i n s tance , a p opu la t ion w i th h i gh p ropo r t ion s o fa du l t d ea th s m ay , a s sum ing t h a t t h e s amp le i sn o t b i a sed a nd t h a t t h ep opu la t ion i ss t ab le , b ee xpe r ienc ing l ow s ub -adu l t m o r ta l i ty . T he m o r ta l i ty p a t te rn , h oweve r , c an b e a na lyzed f u r the r b y a s sum ing t h a tt h ep a t te rn m igh tb e m is lead ing . C hange s i n f e r t i l i ty r a te s a nd m ig r a t ion m ay a c coun t f o r t h e r e su l t s . H ypo the t ica li n c rea se so rd e c rea se si nt h e f e r t i l i ty r a te o ft h ep opu la t ion u nde rc on s ide ra t ion

I ntroduc t ion T he f i e ld o f p a laeodemog raphy i sr e la t ive ly n ew . I th a sd eve loped s ub sequen tt ot h ea dvance s m ade b y s k e le ta l b i o log i s t s a nd t h e g r ow ing a wa r ene s s t h a t h uman s k e le ta l r ema in s a r e v i ta l a r chaeo log ica lr e sou r ce s . P a la e ,odemog raphy h a s t h e p o ten t ia l t op l ay ak ey r o le i nd i s c lo s ing t h e i n t r ica c ie s o fp a s t l i f e a nd d ea th p a t te rn s . T h i s h o ld s e spe c ia l ly t r ue f o r p a t t e rn s b e l ieved t ob e m an if e s ti nm ed ieva lB r i ta in . A l though t h e f i e ld o f d emog raphy h a s b e en u s ed e x ten s ive ly t o r e con s t r uc t m ed i eva l l i f e , m o s t s t ud ie s h ave b een l i m i ted t o w e l l d o cumen ted p e r iod s o f m ed ieva l h i s to ry , t h a t i s , t ot h e l a te r p e r iod s . Many s t ud ie s , f o r i n s tance , r e ly h eav i ly u pon p a r i sh r e co rd s ( Cowg i l l , 1 963 , 1 967 ) . B u t t h e se r e co rd s w e r e n o t m ade c ompu l so ry u n t i l 1 538 ( P a l l i se r , 1 9 79 :112 ) . S im i la r ly ,t h eu s eo ft omb s tone sd i dn o tb e come a p opu la r m ean s o f m a rk ing g r ave s .w i th i n fo rma t ion u n t i l t h e 1 7 th c en tu ry ( J ohn son , 1 9 12 ) ,t h e r eby a l so s e rv ing a s ac on s t r a in t . Wh i le m ano r ia l r e co rd s h ave b een u s ed t or e c r ea te t h e m ed ieva l d emog raph i c s t ru c tu re o fr u ra l v i l lage s ( Raz i , 1 980 ) , t h ey a r e o f l i t t le a s s i s tan ce i n r e con s t ruc t ing t h e d emog raphy o fu r ban c en t re s . M ed ieva lw i l l sa nd t h el a yp o l lt a xh ave a l so b e en u t i l ized a s t o o l s f o r u nde r s t and ing m ed ieva l l i f e ( Ba r t le t t , 1 9 53 ; H ea th , 1 984 ) b u t " w i l l -mak ing s e em st oh ave b een ah ab i ta cqu i r ed b yt h en ew ly w ea l thy , m e r chan t s a nd c r a f t smen i np a r t i cu la r , t owa rd s t h e e nd o f t h e 1 4 th c en tu ry a nd o nce a dop ted b y t h em b e came f a sh ionab le " ( Ke rmode , 1 9 82 :10 ) . A p rob lem o f b i a s t h u s a pp ea r s e v iden t i n r e sea r ch c once rn ing m ed ieva ll i f ei nB r i ta in . T he d emog raph i c r e l iance u pon w r i t t en d o cumen ta t ion h a s c on t r ibu t ed l i t t le t owa rd o u r u nde r s tand ing o ft h e l i ve s o ft h e u r ban p oo r a nd t h el i ve so fw omen a nd c h i ld r en . T he g oa lo ft h i s p ape r i s t o e xp lo re t h e w ay s i n w h ich a p a laeodemog r aph i c a na ly s i s c an a s s i s t i nf i l l ing t h e se g ap si no u rk now ledge . P a laeode mograph ic Ana lys is P a laeodemog raph i c e va lua t ion o f s k e le ta l p opu la t ion s h a s b e come a p r ima ry m ean s o f h i gh l igh t ing t h e p r e sence o fs t re s so r s . L i f e t a b le a na ly se s , w h i ch i n c lude t h e c a l cu la t ion o f m o r ta l i ty r a te s , s u rv ivo r sh ip c u rve s a nd l i f e e xpe c tan cy r a te s , l e d r e sea r che r s s u ch a s L a l lo ( 1 973 ) , G r een e t a l ( 1 974 ) , S wed lund a nd

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We i s s ( 1 973 ) , w h ich m igh t e x i s t i f s ubadu l t s c on t r ibu t e t o l e s s t h an 3 0% o f t h e t o ta l p opu la t ion . T he s u rv ivo r sh ip c u rve ( I x )c an a l so s e rve a sa v i sua l c l ue t op a laeodemog raph i c p a t te rns w i th in a nd b e twe en p opu la t ion s . I t i s c on s t ruc ted b y s ub t ra c t ing t h e p ropo r t ion d ead a t e a ch a ge i n te rva lf r om 1 00 p e r cen ta l ive a tb i r th . A l though s u rv ivo r sh ip c u rve s d o n o t l e nd t h em se lve s t o s t a t i s t ica l e va lua t ion , r e sea r che r s h ave u s ed t h em t od e t ec t d i f f e r ence s i n mo r ta l i ty p a t te r n s c au sed b y d i f f e r ing c ond i t ion s d u r ing l i f e . A s a n e xamp le , G re en a a l ( 1 974 ) c ompa r e t h e s u rv ivo r sh ip c u rve s o f i n d iv idua l s b u r ied w i th c eme t e ry s up e r s t ruc tu r e s t o i n d iv idua l s b u r ied w i thou ts up e r s t r uc tu re s . T he p a t t e rn s ,t h ea u tho r s c on t end , s how t h a t b u r ia l s w i th s up e r s t ru c tu r e s m ay r e f le c ti n d iv idua l so fh i ghe rs t a tu s . L i f e e xpec tancy r a t e s ( e ° x)a r e a l so u s e fu l i n t e r - a nd i n t r a -popu la t ion m ea su r e s . S eve r a l r e sea r che r s h ave e xam ined c hange s i n l i f e e xp e c tancy r a t e s b e tween p opu la t ion s o ve r t i me ( s ee A c sad i a nd N eme ske r i , 1 970 ; R u s se l l , 1 948 ) ; B ro thwe l l , 1 971 ; A nge l , 1 969 a nd We i s s , 1 973 ) . T he se s t ud ie s h ave c ha l lenged t h e r e sea r che r s t o e xp lo r e b i o log ica l a nd c u l tu ra l d i f f e r ence s b e tween t h e p opu la t ion s a s p o s s ib le m eans o f e xp la in ing v a r ia t ion . O the r r e sea r che r s ( La l lo , 1 9 73 ; L a l lo a a l , 1 978 ; G r een e ta l , 1 9 74 ) h ave a s se s sed c hange s i nl i f e -expe c tancy r a t e s w i th in a p opu la t ion o ve rt i me . Y e ta no the ra pp l i ca t ion h a s b e en t owa rd s a s se s s ing d i f f e r ence s i n p a t t e rn s a c co rd ing t o d i f f e rence s i n s t a tu s ( G r een e t a l , 1 9 74 ; A rme lago s a a l , 1 98 1 ) . T he se s t ud ie s s u gge s t t h a t i n c r ea sed s t a tu s b rough t i n c r ea sed l i f e -exp ec tancy r a te sw i th in ap opu la t ion .

w i l lc oncom i tan t ly c hange t h ep a t t e rn o f mo r ta l i ty f o r t h e p opu la t ion b e ing t r ea ted a s ac oho r t . T he m o r ta l i ty p a t te rn m ay i n f a c t b e t o p -heavy n o t b ecau se o f l ow f e r t i l i ty , h i gh m ig ra t ion o f a du l t s i n to t h e c ommun i ty o r h i gh m ig ra t ion o f s ubadu l t s o u t o f t h e c ommun i ty . Moo r e e ta l ( 1 975 ) , J ohan s son a nd H o row i tz ( 1 986 ) a nd H o row i tz a nd A rme lago s ( 1 988 ) s ugge s t t h a t r e sea r che r s b ecome s e n s i t ive t o t h e e f f e c t o f i n s tab i l i ty u pon ap opu la t ion . S en s i t iv i ty o ccu r s w hen t h e r e sea r che r a sk s h ow t h e e f f ec t s o f i n c rea s ing o r d ec r ea s ing f e r t i l i ty r a t e s o f c hang ing m ig ra t ion p a t te rn s w i l l a f f e c t t h e p a leodemog raph i c m ea su re s . T he r e su l t s o ft h e i n i t ia l a s se s smen t c a r r ied o u t u nde r t h e a s sump t ion o f s t ab le p opu la t ion c ond i t ion s , t h e r e fo r e , a r e n o t a dop ted a s s t r ic t i n d ica t ion s o f t h e p opu la t ion ' s d ynam ics . T hey a r e t r ea ted a s t h eb a s i sf o rf u r the re xp lo ra t ion . T he f i na l c r i t ic i sm , t h a t r e su l t s o f l i f e t a b le a na ly se s l a ck c o r r e la te s t ol i v ing p opu la t ion s , h a s b een a dd r e s sed b y G r eene e ta l( 1 986 ) . H e r e t h e a u tho r s s how t h a t t h e i r r e su l t s f r om t h e K u lubna r t i s k e le ta l p opu la t ion a ppea r s im i la r t o s e ve ra ll i v ing p opu la t ion s . I ta ppea r s a tt h i s t i me t h a t p a laeodemog r aphy c an c au t iou s ly b e u s ed t o a s se s s p a t te r n s o f a dap ta t ion a nd c hange i n a r cha eo log ica l p opu la t ion s . T h i s s t udy w i l l u s e l i f e t a b le s t o e xam ine t h e p r e sence o f s t r e s so r s i n t h e S t . H e len -on the - Wa l l s p opu la t ion , f o cu s s ing p r ima r i ly u pon m o r ta l i ty r a t e s , s u rv ivo r sh ip c u rve sa nd l i f e -expe c tancy r a te s . T he f o rmu la t ion o f l i f e t a b le s a l low s f o r a c r i t ica l e xam ina t ion o f ap opu la t ion . L i f e t a b le s c on s i s t o f s e ven c o lumns r e p r e sen t ing t h e a ge i n te rva l ( x ) , t h e a c tua l n umbe r d ead i ne a ch a ge i n te rva l ( d ix ) ,t h ep e r cen tage o fi n d iv idua l sd y ing a ta n a ge i n te rva l o u t o ft h e t o ta l p opu la t ion ( d x ) , t h e p e r cen t l i v ing a t a n a ge i n t e rva l ( I x ) , t h e p robab i l i ty o f d y ing a ta n a ge i n te r va l ( q x ) , t h e n umbe r o f y ea r s l i ved b y t h e t o ta l p opu la t ion b e tween a ge x a nd x + 1 ( Lx ) , a nd t h e c a lcu la ted l i f ee xpe c tancy r a t e a ta n a ge i n te rva l ( e ° x) .L i f e t a b le s t h u s u se b o th t h e mo r ta l i ty r a t e o f t h e p opu la t ion a nd t h e s u rv ivo r sh ip c u rve t oa r r ive a t al i f e e xpe c tancy r a te a nd m ean a ge o f d ea th f o r t h ep opu la t ion . T he a na ly se s o f m o r ta l i ty r a te s ( d x ) a r e p a r t icu la r ly u s e f u l t o t h e s t udy o f p a s t h uman p opu la t ion s . S i n ce m o r ta l i ty c an b e v i ewed a s t r ue m a ladap ta t ion , a ge i n te rva l s c on ta in ing h i gh n umbe r s o fd ea th s a r e e sp ec ia l ly i mpo r tan t t ot h e r e sea r che r , f o r t h ey c an e l uc ida te a ge i n te rva l s o f g r ea t s t re s s t om embe r s o f t h e p opu la t ion o rc an h i gh l igh t t h e p r e sence o fs amp le b i a s . B ro thwe l l ( 1 971 ) , We i s s ( 1 973 ) a nd B u ik s t ra ( 1 981 ) u s e m o r ta l i ty r a te a s se s smen t s a s g u ide s t o s amp le b i a s . Mo s t c ommon i st h e p o ten t ia l f o r i n f an t u nde r -enume ra t ion , w h ich c an b e d e te c ted b y d i v id ing t h e n umb e r o f i n f an t s ( 0 1 y ea r o l d ) b y t h e n umbe r o f s ubadu l t s ( 1 -20 y ea r s o l d ) i nt h e p opu la t ion ( Bro thwe l l , 1 971) , o r , a c co rd ing t o

T he S amp le T he m ed ieva l c eme t e ry o f S t . H e len -on the Wa l l s ( c i rca 100 -1550 A .D . ) a tY o rk , E ng land , p r ov ided t h ree c r i t ica l r e qu i s i te s f o r a na ly s i s . F i r s t , i t p r ov ided a n h omogeneou s s k e le ta l s amp le . D o cumen ta ry s ou r ce ss ugge s tt h a tt h eS t . H e len -on the - Wa l l s p a r i sh ione r s w e r e a mong s t t h ep oo r e s t i nt h ec i ty . S econd , a l l a ge s a nd b o th s e xe s w e r e p r e sen t i nt h e s amp le , s ugge s t ing t h a t b l a tan tb u r ia ls e le c t ion c r i te r ia w e r en o tp r a c t iced . L a s t , t h e s amp le s i ze ( N = 1 0 14 ) w a s l a rge e nough t o a l low s t a t is t ica l e va lua t ion o f t h e t r end s .

E xpec ta t ion s P a laeodemog raphy c an s e rve a s am ean s o f a s se s s ing t h e p r e sence o f s t r e s so r s w i th in a p opu la t ion w i thou t r e qu i r ing t h e i s o la t ion o f t h e s p e c i f ic s t r e s so r a t w o rk . U s ing t h i s p r em i se , s e ve ra l p a laeodemog raph ic e xpec ta t ion s c a n b e f o rmu la t ed f o r t h e S t . H e len s p opu la t ion . T he e xpe c ta t ion s a r e c on t ingen t u pon t h e a s sump t ion t h a t s t r e s s w i th in t h e p opu la t ion w i l l b e d e te c t ed i nt h e m o r ta l i ty p a t te rn . I ti sa l so a s sumed t h a t

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r ema in s w e r e a d equa te ly p r e se rved o t he r m ean s o f a ge ing w e r e a l so u s ed . T he d eg ree o f e p iphy sea l u n ion , f o ri n s tance , w a s u sed t oa s se s s t h e a ge o fs ubadu l t s ( McKe rn a nd S t ewa r t , 1 957 ; K rogman , 1 962 ; U b e lake r , 1 978 ) ,w h i le t h ep ub ic s ymphy s i s w a su s ed a sa na ge i n d ica to rf o ra du l t s ( Todd , 1 920 , 1 92 1 ; McKe rn a nd S t ewa r t , 1 957 ; Me ind l e t a l , 1 985 ) . T he l a s t m eans o f a s se s smen t u s ed c r an ia l s u tu r e c l o s ing o n ly w hen o t he r m ean s o f a ge ing w e re d i f f icu l t o r i mpo s s ib le . T he se f o u r t e chn ique s a l lowed f o r t h e e ven tua l s e r ia t ion o ft h e s k e le ta l m a te r ia l b y a ge . T he s e r ia t ion o f t h e p opu la t ion r e l ied u pon t h e d eve lopmen ta l a ge o f t h e i n d iv idua l a nd d i d n o t a s sume t h a t a r cha eo log ica l p opu la t ions c ou ld b e a s s igned a b so lu te a ge s u s ing am ode rn r e f e rence p opu la t ion . S ex w a s a s se s sed u s ing t h ree s epa r a te t e chn ique s , t h e f i r s t r e qu i r ing av i sua l e va lua t ion o f t h e p e lv i s ( B ro thwe l l , 1 981 ) . T he s ubpub ic a ng le a nd s c ia t ic n o t ch w e re u s ed a s p r ima ry d e t e r inan t s o f s e x ( K rogman , 1 946 , 1 962 ; Wa shbu rn , 1 948 ; P hen ice , 1 967 , 1 969 ) . S ex w a s a l so d e te rm ined u s ing f e a tu re s o f t h e mand ib le a nd c r an ium ( Keen , 1 950 ) .

s t r e s so r sd on o tu n i fo rm ly a f f e c te ve ry m embe ro f t h ep opu la t ion , b u t , r a the r , d ep end ing o n t h e t y p e o f s t re s so r , w i l l a f f e c t c e r ta in a ge g r oup s mo r e f r equen t ly t h an o t he r s . T he p r e sence o f m a lnu t r i t ion a nd i n fe c t ion i s e xp ec ted t o c on t r ibu te t o ah i gh s ubadu l tm o r ta l i ty r a t ef o rt h e S t . H e lens p opu la t ion d ue t o t h e p a r t i cu la r s u scep t ib i l i ty o f t h e se a ge g r oup s t o s qua l id e nv i ronmen ta l c ond i t ion s . T he s t r e s so r s , n o d oub t , w i l l h ave p l ayed a n i mpo r tan tr o le i na du l t l i f ea sw e l l , c on tr ibu t ing t oa ne xpec t ed l ow m ean a ge a td ea th a nd l ow l i f e -expe c tancy . D i f f e rence s i n t h e m o r ta l i ty r a t e s b e tween m a le s a nd f ema le s a r e a l so e xpe c t ed . A n i n c r ea sed m o r ta l i ty o f f ema le s d u r ing r e p roduc t ive y ea r s i s e xp e c t ed d ue t o t h e c ompounded p hy s io log ica l d i s r up t ion c au sed b y m a lnu t r i t ion , i n f ec t ion a nd p r egnancy . C owg i l l ( 1 963 ; 1 967 ) h a s s ugge s ted f r om 1 6 th c en tu ry d o cumen ta ry i n fo rma t ion f r om t h e c i ty o f Y o rk t h a t " a l though t h e d a ta i n d i ca te n o s t a t i s t ica l ly s i gn i f ican t d i f f e r ence b e twe en t h e m a le a nd f ema le mo r ta l i ty i n t h e Y o rk d a ta , t h e r e i s a d i f f e rence , p o s s ib ly d ue t o " c hance " , b u t n eve r the les so ccu r r ing a s ar e a lf e a tu r eo ft h es ub p opu la t ion i nq ue s t ion ,t h ef ema le d ea th r a te a ta l t i me s u n t i l t h ee nd o fl i f eb e ing r a the rh i ghe rt h an t h em a le " ( Cowg i l l , 1 967 :6 1 ) . T he p a la eodemog raph ic a na ly s i s o f t h e S t . H e lens p opu la t ion w a s t h e re fo re i n i t ia ted w i th t h r ee g oa ls i nm ind . T he f i r s t g oa l w a s t ou t i l ize p a laeodemog r aph i c a na ly s i s , g i v ing s pe c ia l a ten t ion t ot h e c r i t i c i sm s o ft h e t e chn ique s . T h i s w a s a ch ieved , i np a r t , b y u s ing mu l t ip le a ge ing a nd s e x ing t e chn ique s , w h i ch w e re u sed i no rd e r t oa vo id t h e p o ten t ia l t owa rd s m im i ck ing t h e a ge a td ea th p a t te rn o f as i ng le r e f e r ence p opu la t ion a nd b y e xp lo r ing t h e a pp l icab i l i ty o f t h e s t ab le p opu la t ion t h eo ry i n t h e a na ly s i s a nd t h e S t . H e len -on the - Wa l l s p opu la t ion . T he s e cond g o a l w a s t o u se p a laeodemog r aph i c m e a su re s a s a m ean so ff o rmu la t ing i n f e r ence sa bou tp opu la t ion p ro ce s ses w o rk ing i n m ed ieva l u r ban c en t r e s . T he l a s t g o a l w a s t o a s se s s w he the r t h e p a laeodemog r aph ic p a t t e rn s w e r e c on s i s t en t w i th t h ee xpec ta t ion s .

R e su l ts a nd D iscuss ion T he e xcava t ion o ft h e S t . H e len -on the - Wa l l s c eme te ry y i e lded 1 0 14 d i s c r e te ly b u r ied s k e le ton s . O f t h e se , 6 37 i n d iv idua l s w e r e a s s igned t o a ge -g roup i n t e rva l s , w h ich r a nged f r om o ne -yea r i n t e rva l s u p t ot h e a ge o f 5 ,f i ve y ea r i n te rva l s u p t ot h e a ge o f 2 5 , a nd t e n -yea r i n te rva l su pt ot h em ax imum a ge o f6 5 . As um o f 3 77 s k e le ton s c ou ld o n ly b e a s s igned a s a " s ubadu l t " ( 0 -20 y ea r s o l d ) o r a n " adu l t " ( 20 -65 y ea r s o l d ) d ue t ot h e s e ve r e ly f r agmen ta ry n a tu r e o f t h e s k e le ta l m a t e r ia l . T he se i n d iv idua l s w e r e p l a ced i nt h em o s tr e f ined a ge c a t ego r ie sp o s s ib le . T he f o l low ing p a la eodemog raph ic m ea su r e s w e r e p r o ra ted , u s ing t h e p ropo r t ion d y ing a te a ch a ge g r oup i n t e rva l o ft h o se c apab le o fb e ing a ged a sa r ef l ec t ion o f t h e t o ta l p opu la t ion . T he c l ea r ly a ged i n d iv idua l s w e re u s ed a s ag u ide t o t h e p ropo r t iona te p l a cemen t o f t h e mo re d i f f icu l t i n d iv idua l s .

Me thods a nd T echn iques T he p a laeodemog r aph ic a na ly s i s o f t h e S t . H e lens s ke le ta l p opu l a t ion r e qu i r ed a d e te rm ina t ion o fs e x a nd a ge f o ra l i n d iv idua l s i n t h e p opu la t ion . F ou r t e chn ique s w e r e e mp loyed t o a ge t h e s k e le ta l m a t e r ia l . T he p r ima ry t e chn ique w a s d en ta l a tr i ton r a t e s ( M i le s , 1 963a ; B ro thwe l l , 1 981 ; L ove joy , 1 985 ) a nd d en ta l e r up t ion p a t te rns i n j uven i le s ( S chou r a nd M a s s le r , 1 941 ; H un t a nd G le i se r , 1 955 ; M i le s , 1 963b ; Moo r re e s e ta l , 1 963a , 1 963b ; B ro thwe l l , 1 981 ) . T he se t e chn ique s w e r e c ho sen d ue t ot h e f r equen t ly f r agmen ta ry n a tu r e o f t h e p o s t c r an ia l s k e le ta l m a te r ia l a nd t h e c ompa ra t ive ly g ood c ond i t ion o ft h e d en t i t ion . When t h ep o s t c r an ia l

a )P opu la t ion P a t tern s L i f e t a b le s w e r e f o rmu la ted t o a s se s s t h e m o r ta l i ty p a t te rn a nd l i f e e xpec tancy r a t es o f t h e S t . H e len s p opu la t ion . I t w a s p ropo sed t h a t s t re s so r s a s so c ia ted w i th t h e S t . H e len s s o c io e conom ic e nv i ronmen t w ou ld c on t r ibu te t o h i gh i n f an t mo r ta l i ty r a t e s a nd a g ene ra l ly l ow l i f e e xpe c tancy r a t ef o rt h i sp opu la t ion . T he l i f e t a b le f o r t h e S t . H e len s p opu la t ion c an b e f o und i nT ab le 7 . 1 . T he mo r ta l i ty r a te , c o lumn d x , w h ich i sg r aph ica l ly r e p r e sen ted i n F i gu r e 7 . 1 , i n d ica t e s t h a ta ge -g roup i n te rva l 5 10 , i n w h ich 1 .1% o f a l l d ea th s o ccu r , a ge g r oup

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i n te rva l 2 5 -35 , i nw h ich 2 3% o fa l l d ea th s o c cu r a nd a ge g roup i n te rva l 3 5 -45 , i nw h ich 2 0 .1% o f t h e d ea th s o ccu r , h ave t h e g r ea te s tm o r ta l i ty r a t e s o fa ny a ge g roup i n t e rva l s . T he c a l cu la t ion o fl i f e e xpe c tancy r a te s , c o lumn e ° x,i n d ica te s t h a t a t b i r th a n i n d iv idua l f r om S t . H e len s c ou ld e xpe c t t ol i ve 2 9 .2 m o re y ea r s . T he m o r ta l i ty p a t te rn a nd t h e l i f e e xp ec tancy r a te a r e p e rp lex ing i n l i gh t o f t h e i n i t ia l p r ed i c t ion s . F o r i n s tance , t h e S t . H e len s p opu la t ion w a s e xpe c ted t o d i sp lay h i gh i n f a n t m o r ta l i ty r a te s a s ar ef le c t ion o ft h e s u scep t ib i l i ty o f c h i ld r en t o t h e e f f ec t s o f m a lnu t r i t ion a nd i n f e c t ion . H oweve r , t h e mo r ta l i ty r a t e s a r e n o t o n ly l ow f o r a l y oung s ubadu l t a ge -g roup i n te rva l s u p t ot h e a ge o f f i ve i nc ompa r i son t o o t he r a ge -g roup i n t e rva l s , t h e p e r cen tage o f s ubadu l t s v e r su s a du l t s i n t h e S t . H e len s p opu la t ion i sa l so l ow . When t h e S t . H e len s p opu la t ion i s d i v ided i n to t wo a ge g roup s , s ubadu l t ( a ge s 0 20 ) a nd a du l t ( a ge s 2 0 -65 ) , s ubadu l t sa c coun tf o ro n ly 2 8% o ft h et o ta lg r oup , ap ropo r t ion t h a t i sl ow w hen c ompa r ed t om ode l l i f e t a b le s ( We i s s , 1 973 ) . P e rp lex ing , t o o , i st h e s e em ing ly h i gh l i f e e xp ec tancy r a t e a tb i r th f o r t h eS t .H e len sp opu la t ion . R u s se l l ( 1 948 ) h a s r e con s t ruc t ed l i f e e xpe c tancy p a t te rn s f o r m ed ieva l E ng land u s ing d o cumen ta ry e v idence , t h u s r e c r ea t ing t h e d emog raph ic s t ru c tu r e o f t h e w ea l thy . W i th a m ax imum a ge a td ea th o f9 0 y ea r s , R u s se l l f i nd s t h a t t h e e xpec ta t ion o fl i f e a tb i r th r a nge s f r om a h i gh o f 3 5 .3 y ea r s b e fo re 1 276 , t o al ow o f 1 7 .3 y ea r sd u r ing t h ep l ague y ea r so f 1 346 1375 . T h i s a ve rage s t o al i f e e xpe c tancy o f 2 6 .3 f o r y ea r s b e tween 1 276 a nd 1 450 . We i s s ( 1 973 ) s ugge s t s t h a t t h e " d e f ic i t i n j uven i le s a cknow ledged b y R u s se l lw ou ld t e nd t or a i se t h e mo r ta l i ty r a t e sa nd l owe r t h e l i f e e xp ec tanc ie s s t i l l f u r the r " ( We i s s , 1 973 :42 ) , i n c rea s ing t h e d i spa r i ty b e tween t h e S t . H e len l i f e e xpec tancy a t b i r th a nd t h o se c a lcu la t ed b yR u s se l l . I ti sw hen t h e S t . H e len s l i f e t a b le d a ta a r e c ompa red t o mod e l l i f e t a b le s t h a t t h e d i sc r epanc ie s a nd p a t te rns c an b e a s se s sed . C ompa r i son sb e tween t h eS t . H e len sl i f et a b le a nd We i s s ' ( 1 973 ) m ode l l i f e t a b le s h i gh l igh t t h r ee i n con s i s tenc ie s w i th in t h e S t . H e len s l i f e t a b le d a ta . We i s s a s se r t s t h a t 4 9 o f t h e 7 2 m ode l l i f e t a b le s f o l low a p r ed i c tab le mo r ta l i ty p a t ter n . F i r s t , t h ey i n d i ca te t h a t t h e p robab i l i ty o f d y ing ( c o lumn " qx " ) w i th in t h e f i r s t y ea r o f l i f e i s g r ea te r t h an t h e p robab i l i ty o fd y ing b e tween t h e a ge s o f 1 5 -20 . S e cond , t h e p robab i l i ty o f d y ing b e tween t h e a ge s o f 1 5 -20 i sg r ea te r t h an d y ing b e twe en t h e a ge s o f 1 0 -15 . A nd l a s t , t h a t t h e p ropo r t ion o fj uven i le d ea th s i nt h e p opu la t ion i s g r ea te r t h an 3 0% o f a l d ea th s . C a lcu la ted l i f e t a b le su nab le t os a t i s f y t h e se c ond i t ion s s hou ld b e " c on s id e r ed a s u n l ike ly t o mode l r e a l s t a t iona ry c ond i t ion s a nd s hou ld b e u s ed o n ly i ft h e r e i s c ompe l l ing e v idence t od os o " ( We i s s , 1 973 :47 ) .

T he r e su l t s o ft h e S t . H e lens l i f e t a b le f a i l t o s a t i s f y a ny o ft h e se c r i te r ia . A s c o lumn " qx " o f T ab le 7 . 1 i n d i ca te s , t h ep robab i l i ty o fd y ing a tt h e a ge g r oup i n te rva l 0 1 i s . 0 26 , w h ich i s c on s ide rab ly l owe rt h an the p robab i l i ty o fd y ing a t t h e a ge g r oup i n te rva l 1 5 -20 ( q x = . 0 78 ) . S im i la r ly , t h e p robab i l i ty o f d y ing a ta ge 1 0 -15 ( q x =. 0 81 )i sh i ghe rt h an t h ep robab i l i ty o fd y ing b e twe en t h e a ge s o f 1 5 -20 . T he p ropo r t ion o f s ubadu l t d ea th s i n t h e S t . H e len s p opu la t ion ( 2 8% ) a l so s ugge s t s t h a t t h e p opu la t ion m ay n o t b es t ab le . S i n ce i ta pp ea r s f r om t h e c ompa r i son o f t h e S t . H e len s l i f e t a b le t o We i s s ' mode l l i f e t a b le s t h a t S t . H e len s c anno t b e t r ea ted a s as t ab le p opu la t ion , i tb e come s o bv ious t h a t t r ea t ing t h e S t . H e len s p opu la t ion ' s mo r ta l i ty p a t te r n a nd l i f e e xpe c tancy r a te a tb i r th a sr e a l is t ic i su n ju s t if ied . H oweve r , i td o e s n o t i mp ly t h e r e su l t ing p a t te rn s a r e u s e le s s t owa rd s t h e u nde r s tand ing o f t h e d emog raph i cp r o ce s se so ccu r r ing i nt h i sm ed ieva l p opu la t ion . S eve ra l c au se s f o r t h e p ropo r t iona te ly l ow n umb e r so fs ubadu l t si nt h e S t . H e len sp opu la t ion w e re t e s ted . T he f i r s tp ropo sed c au se o fb i a s w a s t h e p r e sen ce o f d i f f e r en t ia l s ke le ta l p r e se rva t ion a c co rd ing t o a ge . T h i s p ropo s i t ion w a s b a sed u pon t h e r e cogn i t ion t h a t s ubadu l t s ke le ta l m a t e r ia l i sm o r e f r ag i le t h an a du l t m a te r ia l a nd l e s s l i ke ly t o b e p r e se rved . A n a r chaeo log i s t t h e re fo r e h a s ag r ea te r c hance o f e xhum ing a n o l de r -aged i n d iv idua l t h an a y ounge r -aged i n d iv idua l . H oweve r , t h er e su l t s o ft h e S t . H e len s e xcava t ion d o n o t s uppo r t t h is c on ten t ion . T he a ge -g roup i n t e rva l 5 10 , f o r i n s tance , d i sp lay s a g r ea te r n umb e r o fi n d iv idua l s t h an t h e a ge -g roup i n te rva l p r e ced ing i t . T h i s r e su l t s ugge s t s t h a t p r e se rva t ion i sn o t m e r e ly a f unc t ion o f a ge o r b one s i ze . T he s e cond p ropo sed c ause f o r s ubadu l t s k e le ta l b i a s c on s ide r ed d i f f e r en t ia l b u r ia l p r ac t ice s a s af u nc t ion o f a ge . T h i s h ypo the s is w a s b a sed u pon t h e C h r is t ian p r a c t ice o f e xc lud ing u nbap t ized i n f an t s f r om g r aveya rd b u r ia l o ri s o la t ing t h e se b u r ia ls o n t h e n o r th s i de o f t h e g r aveya rd o r c hu r ch s t ruc tu r e ( J ohnson , 1 9 12 ) . My r c ( 1 868 ) , h oweve r , s ugge s t s t h a t m ed ieva l c hu r chmen w e r e c once rned a bou t i n f an t s d y ing b e fo r eb ap t i sm c ou ld b e p e r fo rmed . S ub sequen t ly , m idw ive s a nd o the r l a ymen w e re t a ugh t h ow t ob ap t i ze n ewbor n i n f an t s b e l ieved t o o w eak t os u r v ive . B e r ry a nd S cho f ie ld ( 1 971) , i nt h e i r s t udy o f a ge a tb ap t i sm i np r e indu s t r ia l E ng land , h ave c onc luded t h a t b ap t i sms b e tween t h e p e r iod o f 1 650 1700 t o ok p la ce w i th in t h e f i r s t m on th o fl i f e . T he r e fo r e , i nt h e a na ly s i s o f b u r ia l s i nt h e S t . H e len s p opu la t ion i n f an t s a ged 0 1 w ou ld b eu s ed t ot e s t w he the rb ap t i sm p l ayed ar o le i nm o r tua ry b ehav iou r . I f t h e S t . H e len -on the - Wa l l s c ommun i ty c ho s e t oi n c lude b u ts e g rega t e u nbap t ized i n f an t s , o ne a r ea w ou ld d i sp lay a d i sp ropo r t iona te ly h i ghe r n umb e r o f i n f an t b u r ia ls . T o t e s t f o r t h i s

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r e nde r ing t h e m ean a ge a td ea th o f2 9 .2 y ea r so l d f a rt o oh i gh . T he r e su l t s m ay b e f u r the r e xa ce rba t ed b y u nequa l m ig ra t ion p a t te rn s w i th in Y o rk 's p a r i she s . I tm u s t b e r ememb e r ed t h a t Y o rk ' s p a r i she s d i f f e r ed s o c io -e conom i ca l ly f r om o ne a no the r . T he S t . H e len -on the - Wals p a r i sh , a c co rd ing t ot h eL ay P o l l T ax R e tu rn s ,a ppea r s t o h ave b een o n t h e l ow e nd o f t h e s ca le . T he p a r i sh ione r s w e r e c apab le o f p ay ing o n ly t h e s ma l le s t a moun t o f t i he s w hen c ompa red t ot h e p aymen t s p a id b y m embe r s o f o t he r p a r i she s i n t h e c i ty ' s c en t r e . T he re fo re , w hen u s ing d o cumen ta ry e v idence t o a s se s s t h e a moun t o f i mm ig ra t ion i n to t h e c i ty a nd t h e a s se s sed w ea l th o f ap a r i sh ,t h er e sea r che r mu s tr e cogn ize t h a tt h e r e co rd s w e r e o f t en b i a sed i n f a vou r o f Y o rk ' s m o r e p ro sp e rou s c i t izen s . M ig ran t s , c om ing t o t h e c i ty i n s e a r ch o f w o rk , m ay h ave b een f r equ en t ly o m i t ted f r om t h e P o l l T ax r e co rd s a nd m ay h ave c on s t i tu ted ag r ea t e r p e r cen tage o f t h e p oo r e r p a r i she s t h an o f t h e w ea l th ie r p a r i she s , w h ich a r e w e l l r e p r e sen ted i n d ocumen ta ry e v idence . F u r the r e xam ina t ion o ft h e m ig ra t ion p a t te rn s i n to Y o rk i se s sen t ia l t owa rd s t h e u nde r s tand ing o ft h e c i ty ' s p opu la t ion p ro ce s se s . T he m ig ra t ion o f a du l t s i n to a c ommun i ty n o t o n ly s kew s t h e p e r cen tage s b e tween j u ven i le s a nd a du l t s i n t h e p opu la t ion , b u t m ay h ave a p ro found e f f e c t o n f e r t i l i ty p a t te rn s . B o th R u s se l l ( 1 948 ) a nd G o ldbe rg ( 1 986 ) h ave s ugge s t ed t h a t an eed f o r p e r sona l e conom ic s e cu r i ty d e layed t h e a ge o f m a r r iage i nu rban c en t re s . T h i s d e lay m ay h ave i nf luenced f e r t i l i ty r a t e s w i th in t h e c i ty . I ft h e a ge o f m a r r iage w a s d e layed , t h e n umbe r o f o f f sp r ing m ay h ave b een r e duced . F o r t h e S t . H e len s p opu la t ion t h i s w ou ld h ave c on t r ibu ted t o al ow g r ow th r a te ,o r an ega t ive g r ow th r a te i ft h e b i r th r a te o ft h e p opu la t ion c ou ld n o t b a lance t h e m o r ta l i ty r a te . T he p r e sence o fl ow o rn ega t ive f e r t i l i ty r a te s a l so c on t rad ic t s t h e l i f e t a b le a s sump t ion o f p opu la t ion s t a t iona r i ty . T ak ing h e ed o f t h e d i s cu s s ion so f Moo r e e ta l( 1975 ) ,J ohan s son a nd H o row i tz ( 1 986 ) a nd H o row i tz a nd A rme lago s ( 1 988 ) ,c once rn ing t h ee f f ec t so fc hang ing g row th r a t e s o n l i f e t a b le r e su l t s , t h e r e spon se o f t h e S t . H e len s p opu la t ion l i f e t a b le t o l ow o r n ega t ive f e r t i l i ty r a te s w a s e xp lo red . A cco rd ing t o t h e se a u tho r s , i n as t ab le p opu la t ion t h e m ean a ge a t d ea th w i l l d ec rea se a s t h e f e r t i l i ty r a te o f t h e p opu la t ion i n c rea se s . When t h e S t . H e len s p opu la t ion i s h ypo the t ica l ly t r ea ted a s s t ab le ( m ig ra t ion i n to t h e c ommun i ty e qua l s t h e m ig ra t ion r a te o u t o f t h e c ommun i ty o r n o m ig ra t ion i so ccu r r ing i ne i the rd i re c t ion ) a nd t h e r e la t ion sh ip b e twe en t h e S t . H e len s m ean a ge a t d ea th a nd l i f ee xpe c tancy r a te i sa l t e r ed t oa s sume a+ 1 /2% r i se i nt h e g r ow th r a te , i ta ppea r s t h a t w h i le t h e l i f e e xpec tancy r a te r ema in s c on s tan t , t h e m ean a ge a td ea th w i l l d e c r ea se f r om 2 9 .2 y ea r s t oa pp rox ima te ly 2 5 y ea r s . C onve r se ly , i f

o ccu r rence t h e n o r th s i de o f t h e c eme t e ry w a s s e g r ega ted f r om t h e o t he r a r ea s o f t h e c eme te ry . AC h i squa r e t e s t ( Tab le 7 . 2 ) w a s p e r fo rmed t o a s se s s w he the r t h e n o r th s i de h ad a d i sp ropo r t iona te n umb e ro fi n f an t sa ged 0 1 y ea r s o l d c ompa red t o s ubadu l t s 1 20 y ea r s o l d . T he t e s t i n d ica ted t h a t t h e d i f f e r ence s w e r e n o t s t a t i s t ica l ly s i gn i f i can ta tt h e. 0 5 l e ve l . O n t h eo t he rh and ,i fu nbap t ized c h i ld ren w e r e e xc lud ed f r om g r aveya rd b u r ia l t h en t h e d emog raph ic s t ruc tu re o ft h ec eme te ry p opu la t ion w ou ld a ppea r t o h ave a d i sp ropo r t iona te ly l ow n umbe r o f i n d iv idua l s i n t h e f i r s t a ge -g roup i n t e rva l . T he mo r ta l i ty r a t e f o r t h e S t . H e len s p opu la t ion i n d i ca te s t h a t 2 . 6% o f t h e p opu la t ion d i ed b e fo re t h e a ge o fo ne y ea r , c on t r ibu t ing t oa l ow p robab i l i ty o f d y ing ( q x =. 0 26 ) . Wh i le t h e p robab i l i ty o f d y ing i st h e h i ghe s t o f a l a ge g r oup i n te rva l s b e low t h e i n t e rva l o f 5 10 y ea r s o l d , i ti sv e ry l ow w hen c ompa r ed t oA csad i a nd N eme ske r i ' s ( 1 970 ) a nd We i s s ' ( 1 973 ) mode l l i f e t a b le s , w h ich i n d ica t e t h e p robab i l i ty v a ry ing b e tween . 1t o. 4 . T h i s m ay s ugge s t t h a t t h e l ow p e r cen tage o f s ubadu l t s r e p r e sen ted i n t h e a ge g r oup i n te rva l 0 1 i n t h e S t . H e len s p opu la t ion w a s c aused b y b u r ia l p r a c t i ce s , w h i ch e xc luded t h e b u r ia l o f u nbap t ized c h i ld r en w i th in t h e c onf ine so ft h ec hu r ch c eme te r y . T he r e su l t s o f t h e t e s t s f o r s ubadu l t u nde r e nume ra t ion a r e t h e re fo re r e vea l ing . T hey s u gge s t t h a t t h e b i a s a ga in s t s ubadu l t s i nt h e S t . H e len s p opu la t ion w a s n o t n e ce s sa r i ly d ue t o a r chaeo log ica l p r e se rva t ion b i a se s . R a the r , i t a ppea r s t h a t o t he r p ro ce s se s a r e c on t r ibu t ing t o t h e l ow p e rcen tage o f s ubadu l t s , o ne o f w h i ch m ay b e t h e e xc lu s ion o f u nbap t ized i n f an t s f r om t h ec eme te ry s amp le . H oweve r , t h e r o le o fa du l t s i nt h e S t . H e len s p opu la t ion m u s t n o t b e o ve r looked a t t h i s j unc tu r e . Wh i le a r cha eo log i s t s a nd p hy s ica l a n th ropo log i s t s m u s tb e s e n s i t ive t ot h ee f f e c t s o f p r e se rva t ion b i a se s a ga in s t s ubadu l t s k e le ta l m a te r ia l , t h e p o t en t ia l f o r a du l to ve r -enume ra t ion i n t h e s ke le ta l p opu la t ion m u s t a l so b e c on s ide red . Med ieva l h i s to r ian s s u ch a sR u s se l l ( 1 948 ) p ropo se t h a t u rban c ommun i t ie s w e re m a in ta ined b y h i gh r a te s o fa du l t i mm ig ra t ion . I f t h i s p ropo s i t ion i s t r ue , t h en t h e s k e le ta l p opu la t ion f r om S t . H e len s m ay b e d i sp lay ing a n o ve r -enume ra t ion o fa du l t s , r a the rt h an s imp ly a n u nde r -enume ra t ion o fs ubadu l t s . A s se ss ing t h e n umbe r s o f i n d iv idua l s m ig ra t ing i n to Y o rk i s d i f f icu l t . S o , t o o , i s a s se s s ing t h e p ropo r t ion o f i mm ig ran t s i n to t h e S t . H e len -on the - Wa l l sc ommun i ty . A cco rd ing t o R u s se l l( 1948 ) ,t h eF r eemen ' sR o l l s f o rt h ec i ty o f Y o rk s ugge s tt h a t2 5% o ft h er e co rded i n d iv idua l s w e r e o r ig ina l ly f r om t own s l o ca t ed f i ve t of i f ty m i le s o u t s ide o f Y o rk . T h i s l a rge p ropo r t ion , i f a s sumed t o h ave m ig r a ted t o Y o rk a s a du l t s , w ou ld s kew t h e r e su l t so ft h e l i f ee xpec tancy r a te a tb i r th c a l cu la t ion s f o rt h e S t . H e len sp opu la t ion ,

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t h e S t . H e len s p opu la t ion u nde rwen t p e r iod s w hen r e p lacemen t o f t h e p opu la t ion t h rough b i r ths w a s i mpo s s ib le ,a ss u gge s ted b y h i s to r ian s , t h en t h e m ean a ge a td ea th w ou ld i n c r ea se t o a pp rox ima te ly 3 6 y ea r s o l d , a s sum ing an ega t ive g r ow th r a te o f1% . I t m ay t h e r e fo r e b e p o s s ib e l t h a t t h re e p r o ce s se sw e re t a k ing p l a ce s imu l taneou s ly i nt h e S t . H e len sp opu la t ion ,c au s ing w ha ta pp ea r st ob e a n u nde r -enume ra t ion o f s ubadu l t s i n t h e c ommun i ty a nd a na r t i f ica l ly h i gh l i f ee xpe c tancy r a te a tb i r th . T he f i r s tp ro ce s sm ay h ave b een t h e e xc lu s ion o f u nbap t ized i n f an t s f r om t h e c hu r ch g r aveya rd . T he s e cond m ay h ave b e en ah i gh r a t e o f a du l t i mm ig ra t ion i n to t h e c ommun i ty , s k ew ing t h e c eme te ry s amp le t owa rd s o l d e r i n d iv idua l s . T he l a s t p r o ce s s c on t r ibu t ing t ot h e l i f et a b le r e su l t sm ay h ave b een al ow f e r t i l i ty r a te w i th in t h e c i ty , w h i ch w ou ld r e su l t i n a n i n c r ea sed m ean a ge a td ea th f o rt h ep opu la t ion . T he r e s e ems l i t t le d oub t t h a t t h ec i ty o fY o rk s u f f e r ed f r om m any f l uc tua t ion s i n p opu la t ion s i ze a nd p r opo r t ion . C hange si nY o rk ' se conomy , c oup led w i th r e cu r r ing b ou t s o f p e s t i lence a nd p l ague ,c au sed t h ep opu la t ion o fS t .H e len -on the Wa l l s p a r i sh t o v a ry t h roughou t t h e m ed ieva l c en tu r ie s . P e r iod s o f p opu la t ion i n c r ea se w e r e p unc tua ted w i th e ven t s t h a t c a u sed i mmed ia te d ec rea se s i n s i ze . H oweve r , am o r e g ene ra l p i c tu re o fY o rk ' s p opu la t ion b e tween t h e 1 2 th t o t h e m id 1 6 th c en tu r ie s s how s a n i n c r ea se i ns i ze . Ib e l ieve t h a t t h i s o ve ra l l i n c r ea se w a s d ue , i n p a r t ,t o ah i gh r a te o fa du l tm ig r a t ion i n to t h ec i ty , w h ich , b y c r ea t ing p opu la t ion i n s tab i l i ty , m a sk s t h e p o ten t ia l f o r m o r e s p e c i f ic e va lua t ion s o f f e r t i l i ty r a te s . O bv iou s ly , Ip r opo se t h a t t h e se s ame p r o ce s se s w e r e o c cu r r ing w i th in t h e S t . H e len -on the - Wa l l s p a r i sh a nd c on t r ibu ted t ot h e u nu sua l ly l ow s ubadu l t m o r ta l i ty r a te a nd p r obab i l i ty o f d y ing , a nd t h u s t h e p a r t icu la r ly h i gh l i f ee xp ec tancy r a te a tb i r th i nt h e S t . H e len o n the - Wa l l sp opu la t ion .

i n te rva l c an e xp e c t t ol i ve 2 0 .6 m o r e y ea r s . T he K o lmogo rov -Sm imov t e s t i n d ica te s t h a t t h e d i f f e r ence s b e twe en t h e se t wo p a t ter n s a r e s t a t i s t ica l ly s i gn i f ican t( Tab le 7 . 4 ) . When t h e se r e su l t sa r ec ompa red t om ode ll i f e t a b le s t h e p a t te rn s d i sp layed i n t h e S t . H e len s p opu la t ion b ecome e ven mo r e i n t r igu ing . A c co rd ing t ot h em od e ll i f e t a b les c on s t ru c ted b y C o a le a nd D emeny ( 1 983 ) f ema le s c on s is ten t ly d i sp lay ah i ghe r l i f e e xpe c tancy r a te t h an m a le s . I nt h e S t . H e len sp opu la t ion m a le s c l ea r ly h ave a g r ea te rl i f ee xpec tancy r a te t h an f ema le s ,w h i ch i s g r aph i ca l ly e v iden t i nt h e i rd i f f e ren ts u rv ivo r sh ip c u rve s( F igu r e7 . 3 ) . H e r ei tc an b es e en t h a ta tn o t i me d oe s f ema le s u rv ivo r sh ip s u rpa s s t h a t o f m a le s . T wo e xp lana t ion s f o r t h e g ende r p a t te rn s o f m o r ta l i ty a r e p l au s ib le . T he f i r s t c au se m ay b e t h ep r e sence o f ah i gh n umbe ro fm a te rna ld ea th s . H oweve r , f ema le m o r ta l i ty a ppea r s t op eak a tt h e a ge i n te rva l o f2 5 -35 , al a t e a ge -g roup i n te rva l i f h i s to r ian sa r ec o r r ec ti nt h e i rp r em ise t h a tf ema le s m a r r ied i nt h e i rt e ens ( S haha r , 1 983 ) . T he s e cond c a u se m ay b ed ue t om ig r a t ion p a t t e rn s . Is ugge s t t h a t b o th o f t h e se f a c to r s p l ay a r o le i nt h e S t . H e len sg end e rm o r ta l i ty p a t t e rn s . I n ar e cen t a r t ic le b y G o ldb e rg ( 1 986 ) , t h e t r ad i t iona lv i ew o ff ema le m a r r iage a nd m ig ra t ion p a t t e rn s , w h ich m a in ta in s t h a t m ed ieva l w omen m a r r ied b e fo r e t h e a ge o f t wen ty a nd w e r e u n l ik e ly t ol e ave t h eh ome o ft h e i rp a ren t s u n le s s m a r r ied , i sc ha l lenged . S he a s se r t s t h a t m ed ieva l c i t ie s a nd t own s f r equen t ly s how a h i ghe r p r opo r t ion o f f ema le s t h an m a le s w i th in t h e p opu la t ion . " U s ing p o l l t a x r e turns o f 1 3 77 a sa s o u r ce o n ly 9 0 .5 a du l t m a le s e xc lud ing s e rvan t s a r ef o und t oe ve ry 1 00 w omen i nY o rk ,o n ly 8 6 .1 m a le s i n H u l l a nd a s f ew a s 8 5 .4 i n C a r l is le " ( Go ldbe rg , 1 9 86 :19 ) . T he se l ow s e xr a t io sa r et h e r e su l t o f l a rge n umbe r s o f f ema le s m ig r a t ing t o t h e c i ty a s ar e spon se t ot h e s h if t f r om a r ab le t o p a s to ra l l a nd u s e a nd t h e d ep le t ion o f l a bou r i n t h e c i t ie s a f t e r t h e p l ague . M ig ra t ion , t h e r efo r e , o f f ema le s a s w e l l a s m a le s , p l ayed a r o le i n m a r r iage a nd f e r t i l i ty p a t te rn s : " T he a r gumen t h e r e i st h a t m a r r iage i n t h e l a t e r f o u r t een th a nd f i f te en th c en tu r i e s w a s r e la t ive ly l a te , t h a t i s , f r equen t ly i n t h e m id t wen t ie s a nd v e ry r a re ly i n t h e ' t e en s e ven f o r w omen a nd a l so c ompan iona te i n t h e s e nse t h a t t h e a ge d i f f e r ence b e tween s pouse s w a s n o t v e ry g r ea t . F o r w omen t h e o ppo r tun i ty f o r e mp loymen t ,i ns e rv ice o ra si n dependen tt r ade r s , c r ea ted b y t h e y ea r s o fe conom ic e xpan s ion a nd l a bou r s ho r tage f o l low ing t h e B lack D ea th a lowed t h em t od e lay m a r r iage . . . A bove a l , i t p e rm i t t ed t h em a d eg re e o f i n dependence t h a t y ounge r g i r l s l i v ing a th ome w i th t h e i r p a ren t s c ou ld n o th ave e n joyed ( Go ldbe rg , 1 986 :25 -26 ) . T h i sp a t te rn m ay h ave a f f ec ted t h e S t . H e lens p opu la t ion . When t h e p ropo r t ion o f f ema les t o m a le s i se xam ined , f ema le s c on s t i tu te 5 3 .6% o f t h e a du l t p opu la t ion . T h i s t r end c a nno t b e

b ) G ender P a t terns D i f f e r en t d emog r aph i c p a t t e rn s b a sed u pon g ende r w e re a l so e xpe c t ed t ob e f o und i nt h e S t . H e len s s k e le ta l p opu la t ion . T h i s p ropo s i t ion s t emmed f r om C owg i l l ' s ( 1 963 ; 1 967 ) s u gge s t ion t h a t 1 6 th c en tu ry p a r i sh d ocumen t s f r om t h e c i ty o f Y o rk s howed a h i ghe r f ema le m o r ta l i ty r a te t h an m a le mo r ta l i ty . T he r e su l t so ft h eS t . H e len s i n ve s t iga t ion i n d ica te t h a t m o r ta l i ty r a t e s b y g ende r d i f f e r ( Tab le 7 . 3 a nd F i gu r e 7 . 2 ) . T he f ema le m o r ta l i ty c ur ve a ppea r st or i se t o ap eak a t a ge -g roup i n te rva l2 5 -35 a nd s t ead i ly d ec rea se sa t o l de r a ge -g roup i n te rva ls . T he m a le m o r ta l i ty c u rve r i se s t o ap eak a ta ge -g roup i n t e rva l 3 5 -45 a nd t h en d ec r ea se s s t ead i ly . T he l i f e e xpec tancy r a te u pon r e a ch ing a du l thood ( a ge 2 0 )a l so d i f f e r s b y g ende r . F ema le s r e a ch ing t h e a ge -g roup i n te rva l 2 0 -25 c an b e e xpe c ted t ol i v e 1 8 .3 m o r e y ea r s , w h i le m a le s r e a ch ing t h e s ame a ge -g roup

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a nd f i ve y ea r s o l d , w i th a n a b rup t i n c rea se a tt h e a ge -g roup i n ter va l 5 10 . T he mo r ta l i ty r a te a nd p robab i l i ty o f d y ing p eaked o nce a ga in b e tween t h e a ge -g roup i n te rva l s 2 5 -35 a nd 3 5 -45 . T he l i f e e xpec tancy r a te a tb i r th a ppea red t ob e2 9 .2 y ea r s . When t h e r e su l t s o f t h e S t . H e lens l i f e t a b le s w e re c ompa red t o mode l l i f e t a b le s c on s truc ted b y We i s s ( 1973 ) , t h er e su l t s w e re r e -eva lua ted . I t w a s s ugge s ted t h a t t h ree p roce s ses w e r e r e spon s ib le f o r t h e l i f e t a b le r e su l t s : 1 ) t h a t u nbap t ized i n f an t s w e re e xc luded f r om t h e c eme te ry ; 2 ) t h a t t h e u nde renumer a t ion o f s ubadu l t s w a se xace rba ted b y a n o ve r enume ra t ion o f a du l ts w ho h ad m ig ra ted i n to t h e c ommun i ty ; a nd 3 )t h a t ad e layed a ge a tm a r r iage c ou ld h ave d ampened t h e f e r t i l i ty r a te , t h e reby c au s ing t h e m ean a ge a td ea th f o rt h e S t . H e lens p opu la t ion t o a ppea rh i gh . G ende r p a t te rns w e re a l so e xp lo red . T he r e su l t s i n d ica ted t h a t t h e p eak i nf ema le mo r ta l i ty o ccu r red a t t h e i n te rva l o f 2 5 -35 , w h i le ma le mo r ta l i ty p eaked a t 3 5 -45 . T he se d i f f e rences p roved s t a t i s t ica l ly s i gn if ican t . A lso i n t r igu ing w a s t h e i n c rea sed m a le l i f e e xpec tancy r a te ( 20 .6 y ea r s ) , w h i le f ema le s r e a ch ing a du l thood c ou ld o n ly e xpec t t ol i ve 1 8 .4 mo re y ea r s . T he r e a sons f o r t h i s t r end c ou ld n o t b e e xp la ined b y a s sum ing t h a t t h e S t . H e lens p opu la t ion w as n o t s t ab le . R a the r , i tw a s s ugge s ted t h a t ag r ea te r p ropo r t ion o f f ema le s i n t h e S t . H e lens p opu la t ion w e re m ig ran t s c ompa red t o t h e m a les , a nd t h a t t h e a br up t i n t roduc t ion o f n ew s t res so r s a s soc ia ted w i th m ed ieva l u rban e nv i ronmen t s a nd t h e a dded s t re s so r s a s so c ia ted w i th c h i ldb i r th d ec rea sed t h e l i ke l ihood o ff ema le s l i v ing a sl o ng a sm a le s . T he t h i rd g oa l o ft h is c hap te r w a s t oe xam ine w he the r t h e S t . H e len r e su l t s c omp l ied w i th t h e i n i t ia le xpec ta t ions f o rt h eg roup . I tw a sp ropo sed t h a tt h e S t . H e len sp opu la t ion , b y v i r tue o fi t s l ow s oc io -econom ic p o s i t ion w i th in t h e c i ty o f Y o rk , w ou ld s u f f e r f r om ma lnu t r i t ion a nd i n f ec t ion . T he p r e sence o ft h e se s t re s so r s w ou ld b e i n f e r red f r om h i gh s ubadu l t mo r ta l i ty r a tes a nd ag ene ra l ly l ow l i f e e xpec tancy r a te f o r t h e p opu la t ion . D i f f e rence s b e tween m a le a nd f ema le mo r ta l i ty r a te s a nd l i f e e xpec tancy r a te s w e re a l so e xpe c ted . Women w e re e xpec ted t o s how a g r ea te r mo r ta l i ty a ty ounge ra ge s a nd al owe r l i f e e xpec tancy r a te t h an ma le s . T he r e su l ts o ft h e p a laeodemog raph ic a na ly s i s d i d n o t c omp ly w i th t h e i n i t ia l e xpec ta t ions . A t f i r s t g l ance s ubadu l t mo r ta l i ty a ppea red l ow a nd t h e l i f e e xpec tancy r a te f o r t h e p opu la t ion w a s h i gh w hen c ompa r ed t o o t he r med ieva l p opu la t ion s . H oweve r , t h e r oo t s o f t h e d i sc repanc ie s w e re n o t a s soc ia ted w i th a n i n adequa te a s se s smen t o f t h e S t . H e lens p opu la t ion s oc io -e conom ic c ond i t ion o r w i th a n i n adequa te a s ses smen t o ft h e e f f ec t s o fl ow s t a tus u pon d emog raph ic p roces ses ,b u t ,r a the r , c ou ld b e e xp la ined b y t r ea t ing t h e p opu la t ion a s n on s t ab le . H ence , w h i le m any e xpec ta t ion s w e re n o t m e t b y t h e d a ta , t h e p a leodemograph ic e va lua t ion

a t t r ibu ted t ob i a ses o f s ke le ta l s ex ing t e chn ique s , a s t h e i n co rpo ra t ion o f s ev e ra l s ex ing t e chn ique s c i r cumven ted t h i s t e nden cy n o ted b y We i s s ( 1 973 ) a nd Me ind l e ta l( 1 985 ) . A l though t h e se d i f f e rences w e re n o t f ound t o b e s t a t is t ica l ly s i gn i f ican t u s ing a C h i -Sq ua re t e s t ( Tab le 7 . 5 ) , t h i s p ropo r t ion i s p e rp lex ing a nd m ay i n d ica te t h a t ad i sp ropo r t iona te n umbe r o f w omen w e re m ig ra t ing i n to t h ec ommun i ty . T he p ropo sed p a t ter no ff ema le m ig ra t ion a nd l a te a ge o f m a r r iage a l so s e rve s t o e xp la in t h e p eak i nf ema le mo r ta l i ty a tt h ea ge i n te rva l2 5 -35 . Ad e lay i nm a r r iage a ge c a n b e a s so c ia ted w i th a d e layed a ge o f ma te rna l s t r ess . T he a ge -g roup i n te rva l 2 5 -35 d oes n o ta pp ea rt ob e t o o l a te t ob e c on s idered i n d ica t ive o ft h e p re sence o fm a te rna l s t r es s i f t h e a ve r age a ge o f m a r r iage i n t h e p opu la t ion f a l l s n ea re rt ot h ea ge o f2 0t h an t ot h e a ge o f 1 7 , a nd t h e t r ad i t ion a l v i ew t h a t c h i ldb i r th i n f requen t ly o ccu r red o u t s ide m a r r iage i s a dop ted . H oweve r , a g rea te r p ropo r t ion o f f ema le s m ig ra t ing i n to t h e c i ty o fY o rk a nd ad e layed a ge o fm a r r iage ( a nd t h us m a te rn i ty ) d oe s n o t i ni t se lf e xp la in t h e u nusua l p a t t e rn o f d i f f e r ing l i f e e xpec tancy r a tes b e tween t h e g ende r s . G a l loway ( 1 985 ) h a s s ugge s ted f r om 1 7 th a nd 1 8 th -cen tu ry L ondon d a ta t h a t m id -adu l thood p eak s r e f le c t m ig ra t ion p a t te rns i n to c i t ie s a nd t h e e xpo su re o f n ew i nd iv idua ls t o t h e c i t ie s ' p a thogen ic e nv i ronmen t . P e rhap s ag r e a te r p ropo r t ion o ft h e f ema les i n t h e S t . H e lens p opu la t ion w e re m ig ran ts , w ho , u n l ike t h e g r ea te r p ropo r t ion o f i n d igenous m a les , w e re e xposed t on ew s t re s so r s a l ong s ide t h e s t re s so r s a s so c ia ted w i th c h i ldb i r th . T hu s w hen m ig ra t ion p a t te rn s a r e c oup led w i th m a r r iage p a t te rns f o r t h e c i ty o f Y o rk , t h e d i f f e rence b e tween f ema le a nd ma le mo r ta l i ty p a t te rns may b e e xp la ined b y a dd ing m a te r na l s t re s se s t o t h e o the r s t re s ses e ncoun te red b y w omen r each ing t h ea ge o f2 5 .

C onc lus ion T h i s c hap te r h ad t h r ee g oa ls . T he f i r s t g oa l w a s t oc i r cumven t s ome o ft h e c r i t ic ism a imed a t d i sc r ed i t ing t h e u se o f p a la eodemog r aphy . T he u s e o f mu l t ip le a ge ing t e chn ique s a nd s e r ia t ion r e nde red t h e r ep l ica t ion o f as i ng le r e f e r ence p opu la t ion u n l ike ly . T he s e r ia t ion o ft h e s ke le ta l m a te r ia l p rov ided a n a s se s smen t o f t h e d eve lopmen ta l a ge o f i n d iv idua l s i n t h e p opu la t ion r a the r t h an t h e a b so lu te a ge , t h u s m in im iz ing t h e t e ndency t owa rd s " m im ick ing " r e f e rence p opu la t ion s a s s ugge s ted b y B o cque t A ppe la nd Ma s se t ( 1982 ; 1 985 ) . T he s e cond g oa l w a s t o u se p a laeodemog r aph ic m ea su r e s t o m ake i n f e rences a bou t p a s t mo r ta l i ty a nd mo rb id i ty p a t te rns . L i f e t a b le s w e re c ons t ruc ted f o r t h i s p u rpo se . T he r e su l ts i nd ica ted t h a t t h e S t . H e len s p opu la t ion d i sp layed a l ow mo r ta l i ty r a te a nd a l ow p robab i l i ty o fd y ing f o r i n d iv idua ls b e tween b i r th

7 3

a nd P a t ronage i n t h e F if teen th C en tur y . e d . B . D ob son . p p .G louce s te r : A lan S u t ton P re s s . H o row i tz , S . a nd A rme lagos , G .J . ( 1988 ) . O n g ene ra t ing b i r th r a te s f r om h uman s ke le ta l r ema in s . Am .. 1 .P hy s .A n th ropo l . 7 6 :183 -188 . H owe l l , N . ( 1 982 ) . V i l lage c ompos i t ion i mp l ied b y a p a leodemog raph ic l i f e t a b le : t h e L ibb en s i t e . Am .J . P hys .A n th ropo l .5 9 :263 -70 . H un t , E .E . a nd G le i se r , I .( 1955 ) . T he e s t ima t ion o f a ge a nd s ex o fp reado le scen tc h i ld ren f r om b one s a nd t e e th . Am .J .P hys .A n thropo l . / 3 :479 -88 . J ohan s son , S . a nd H o row i tz , S . ( 1986 ) . E s t ima t ing m o r ta l i ty i n s k e le ta l p opu la t ions : i nf luences o f g row th r a te o n t h e i n te rp re ta t ion o fl e ve l s a nd t r end s d u r ing t h e t r an s i t ion t o a g r icu l tu re . Am . J . P hy s . A n thropo l . 7 1 :233 -50 . J ohn son , W . ( 1 9 12 ) . B yways i n B r i t i sh A rchaeo logy . C amb r idge : C amb r idge U n ive r s i ty P re s s . K een , J .A . ( 1 950 ) . As t udy o ft h e d i f f e rence s b e tween m a le a nd f ema le s ku l l s . Am . J . P hy s . A n thropo l . 8 : 65 -79 . K ennedy , K .A . ( 1984 ) . G row th , n u t r i t ion a nd p a tho logy i nc hang ing p a leodemog raph ic s e t t ing s i n S ou th A s ia . I n P a leopa tho logy a t t h e O r ig ins o f A g r icu l tu re .e d s . M . C ohen a nd G .J . A rme lago s . p p . O r lando : A cad em ic P re s s . K e rmode , J .( 1 982 ) . T he m e r chan t s o f t h re e n o r ther n E ng l i sh t owns . I nP rof e s s ion , V oca t ion a nd C u l ture i n L a te r Med ieva l E ng land . e d . C . C lough . p p . L ive rpoo l : L ive rpoo lU n ive r s i ty P re s s . K rogman , W .M . ( 1946 ) . T he s ke le ton i n f o ren s ic m ed ic ine . P roc .I n s t . Med . 1 6 :154 . K rogman , W .M . ( 1962 ) . T he Human S ke le ton i n F o ren s ic M ed ic ine . S p r ingf ie ld , I l ino i s : C .C . T homa s . L a l l, J .W . ( 1 973 ) . T he S ke le ta l B io logy o f T hree P reh i s to r ic Ame r ican I nd ian S oc ie t ie sf rom D ick son Mound s . P h .D . d i sse r ta t ion , U n ive r s i ty o f Ma s sachuse t t s , Amhe r s t . L a i b, J .W . , A rme lago s , G . J . & R o se , J .C . ( 1978 ) . P a leoep idem io logy o f i n f ec t ious d i sea se i n t h e D ick son Mound s p opu la t ion . Med . C o l l . V i rg in ia Q . 1 4 :17 -23 . L ove joy , C .O . ( 1 985 ) . D en ta l w ea r i n t h e L ibb en p opu la t ion : i t s f u nc t iona l p a t te rn a nd r o le i nt h e a ge d e te rm ina t ion o f a du l t s ke le ta l a ge a td ea th . Am . J . P hys .A n th ropo l .6 8 :47 -56 . McKer n , T .W . a nd S t ewa r t , T .D . ( 1957 ) . S ke le ta l a ge c hange s i ny oung Ame r ican m a les . T echn i ca lR epor t E P -45 . N a t ick , MA : Q ua r terma s te r R e sea rch a nd D eve lopmen t C en te r , E nv i ronmen ta l P ro tec t ion R e sea rch D i v i s ion . Me ind l , R . , L ove joy , C .O . , Men sfo r th , R . & C a r lo s , L .D . ( 1 985 ) . A ccu racy a nd d i rec t ion o fe r ror i n t h e s e x ing o f t h e s ke le ton : i mp l ica t ion s o f p a leodemog raphy . Am .J .P hys .A n th ropo l .6 8 :79 -85 . M i le s , A .E .W . ( 1963a ) . T he d en t i t ion i n t h e a s se s smen t o fi n d iv idua l a ge i n s ke le ta l m a t er ia l . I n D en ta lA n th ropo logy . e d .D . B ro thwe l l . p p .19 1-209 . O xf o rd : P e r gamon P re s s . M i le s , A .E .W . ( 1963b ) . D en t i t ion i n t h e e s t ima t ion o f a ge . J . Den t .R e s .S upp lemen t( 1 )42 :255 -63 . Moo r e , J .A . , S wed lund , A . & A rme lago s , G .J . ( 1975 ) . T he u se o f l i f e t a b le s i n p a leodemog raphy . Am . A n t iq . Mem .3 0 :57 -70 . Moo r ee s , C . , F ann ing , E . & H un t , E . ( 1963a ) . A ge v a r ia t ion a nd f o rma t ion s t ages f o r t e n p e rmanen t t e e th . J .D en t .R es .4 2 :1490 -1502 .

o f t he S t . He len -on the Wa l ls s ke le ta l ma ter ia l p rov ided i n s igh t i n to t he b iases a nd p rocesses a t work w i th in t he c i ty o f Yo rk .

R ef erence s A c sad i , G .Y . a nd N eme ske r i , J . ( 1970 ) . H is to ry o f Human L if e S pan a nd Mo r ta l i ty . B udape s t : Hunga r ian A cad em ic S oc ie ty . A nge l ,L . ( 1969 ) . T he b a se so fp a leodemog raphy . Am . J .P hy s .A n thropo l .3 0 :427 -38 . A rme lago s , G . J . , G oodman , A .H . & B icke r ton , S . ( 1980 ) . D e te rm in ing n u t r i t iona l a nd i n f ec t iou s d i sea se s t re s s i n p reh i s to r ic p opu la t ion s ( a b s t rac t ) . Am .J .P hys .A n thropo l .5 2 :210 . A rme lago s , G .J . , J a cob s , K .H . & Ma r t in , D .L . ( 198 1) . D ea th a nd d emog raphy i n p r eh i s to r ic S udane se N ub ia . I n Mo r ta l i ty a nd I mmo r ta l i ty : T he A n th ropo logy a nd A rchaeo logy o fD ea th . e d s . S .C . H umph rey sa nd H .K ing . L ondon : A cad em ic P r e s s . B a r t le t t , N . ( 1953 ) . T he L ay P o l l T ax R e tu rn s f o r t h e c i ty o f Y o rk i n 1 381 . E as t R id ing H i s t . T ran s . ( o f fp r in t ) . B e r ry , B .M . a nd S chof ie ld , R .S . ( 1971) . A ge a t b ap t i sm i n p r e indu s t r ia l E ng land . P op . S t ud ie s 2 5 :453 -63 . B ocque t -Appe l , J .a nd Ma s se t t , C . ( 1982 ) . F a rewe l l t o p a leodemog raphy . J . Hum . E vo l .2 5 :32 1-33 . B ocque t -Appe l , J .a nd Ma sse t t , C . ( 1985 ) . Ma t t e r s o f momen t . J .H um . E vo l .1 4 :107 -111 . B ro thwel , D . ( 1971 ) . P a laeod emog raphy . I n B io log ica l A spec ts o fD emography . e d . W . B ra s s . p p . 11-30 . L ondon : T ay lo ra nd F ranc i s . B ro thwe l l , D . ( 1981) . D igg ing U pB one s . 3 rd e d i t ion . L ondon : B r i t i sh Mu seum ( Na tu ra lH i s to ry ) . B u ik s t ra , J . ( 1981 ) . Mo r tua ry p ra c t ic e s , p a leodemog raphy a nd p a leopa tho logy : ac a se s t udy f r om t h e K o s t e r s i t e ( I l l ino i s ) . I nT he A rchaeo logy o f D ea th e d s . R . C hapman , I .K inne s , &K . R and sbo rg . p p . 1 23 -32 . C amb r idge : C amb r idge U n ive r s i ty P re s s . B u ik s t ra , J . a nd K on ig sbe rg , L . ( 1 985 ) . P a leodemog raphy : c r i t ique s a nd c on t rove r s ie s . Am . A n th ropo l .8 7 :316 -33 . C oa le , A . J . a nd D emeny , P . ( 1983 ) . R eg iona l Mode l L if e T ab l es a nd S tab l e P opu la t ions 2 nd e d i t ion . O r lando : A cadem ic P re ss . C owg i l l , U . ( 1967 ) . L if e a nd d ea th i n t h e s i x teen th c en tu ry i nt h ec i ty o fY o rk . P op .S tud ie s2 / :53 -62 . C owg i l l , U . a nd H u tch in son , G .E . ( 1963 ) . D if f e ren t ia l mor ta l i ty a mong t h e s exe s i n c h i ldhood a nd i t s p o s s ib le s i gn if icanc e i nh uman e vo lu t ion . P roc .N a t l . A cad .S c i .4 9 :425 -29 . G a l loway , P .R . ( 1985 ) . A nnua l v a r ia t ion i nd ea th b y a ge , d ea th b y c au se , p r ice s , a nd w ea the r i n L ondon 1 670 -1830 . P op .S tud ie s3 9 :487 -505 . G o ldbe rg , P . J .P . ( 1986 ) . F ema le l a bou r , s e r v ice a nd m a r r iage i n t h e l a te m ed iaeva l u rban n o r th . No r th . H i s t . 1 2 :18 -38 . G reen , S , G reen , S . & A rme lago s , G . J . ( 1974 ) . S e t t lemen t a nd mo r ta l i ty o f t h e C h r i s t ian S i te ( 1050 A .D . -1300 A .D . ) o f Me ina r t i ( Sudan ) . J . Hum . E vo l . 3 :297 -316 . G reene , D .L . , V an G e r ven , D .P . & A rme lago s , G . J . ( 1986 ) . L i f e a nd d ea th i na nc ien tp opu la t ion s : b one s o f c on ten t ion i n p a leodemog raphy . Hum . E vo l . 1 : 193 -207 . H ea th , P .( 1984 ) . U rban p i e ty i nt h e l a te r m idd le a ge s : t h e e v idence o f Hu l l w i l l s . I n T he C hu rch , P o l i t ic s

7 4

S chou r ,I .a nd Ma s s le r , M . ( 194 1) . T he d eve lopmen to f t h e h uman d en t i t ion . J . Am . D en t . A s soc . 2 8 :1153 6 0 . S haha r , S . ( 1983 ) . T he F our th E s ta te : AH i s to ry o f Women i nt h e M idd l e A ges . T ran s . C . G a la i . N ew Y o rk : Me thuen . S wed lund , A . a nd A rme lago s , G . J . ( 1 976 ) . D emog raph ic A n th ropo logy . D ubuque , I owa : Wm . C .B rown . T odd , T .W . ( 1 920 ) . A ge c hange s i nt h e p ub ic b one : I . t h em a le w h i te p ub i s . Am .J .P hys .A n th ropo l .3 :285 34 . T odd , T .W . ( 1 921 ) . A ge c hange s i n t h e p ub ic b one . Am .J .P hys .A n thropo l .4 : 1-70 . U b e lake r , D .H . ( 1 978 ) . H uman S ke le ta l R ema ins . C h icago : A ld ine P r e s s . V an G e rven , D .P . a nd A rme lago s , G . J . ( 1983 ) . " F a rewe l lt op a leodemog raphy "? R umo r so fi t sd ea th h ave b een g r ea t ly e xagge ra ted . J . H um . E vo l . / 2 :353 -60 . Wa shbu rn , S .L . ( 1 948 ) . S ex d i f f e renc e s i n t h e p ub ic b one . Am .J .P hys .A n th ropo l .6 : 199 -208 . We i s s , K .M . ( 1 973 ) . D emog raph ic m ode l s f o r a n th ropo logy . Am .A n t iq .3 8 ,P a r t2 , Memo i r2 7 .

Moo rees , C . , F ann ing , E . & H un t , E . ( 1 963b ) . F o rma t ion a nd r e so rp t ion o ft h ree d ec iduou s t e e th i n c h i ld ren . Am .J .P hys .A n thropo l .2 1 :205 -213 . My rc ,J .( 1868 ) . I nI n s t ruc t ion sf o rP a r i sh P r ie s t s . e d . E . P eacock , f r om C o t ton M s . a nd C laud iu s AI . L ondon : T rubne ra nd C o . P a lkov ich , A .M . ( 1 980 ) . P ueb lo P opu la t ion a nd S oc ie ty : A r royo Hondo S ke le ta l a nd Mo r tua ry R ema ins . A r royo H ondo A rchaeo log ica l S e r ie s , V o l .3 . P a l l i se r , D .M . ( 1979 ) . T udo r Y o rk . O xf o rd : O x fo rd U n ive r s i ty P re s s . P hen ice , T .W . ( 1967 ) . A n ew ly d eve loped v i sua l m e thod o f s e x ing t h e o s p ub i s . Am . J . P hys . A n thropo l .3 0 :297 -302 . P hen ic e , T .W . ( 1 969 ) . A n a na ly s i s o f t h e h uman s k e le ta l ma te r ia lf r om b u r ia lm ound si nN o r th C en t ra l K an sa s . U n iver s i ty o f K an sa s P ub l ica t ion i n A n thropo logy ,N o . 1 . L aw rence ,K S . R az i , Z . ( 1980 ) . L if e , Ma r r iage a nd D ea th i n a Med ieva lP a r ish . C amb r idge : C amb r idge U n ive r s i ty P re s s . R u s se l l , J .C . ( 1948 ) . B r i t i sh Med ieva l P opu la t ion . A lbuque rque : U n ive r s i ty o fN ew Mex ico P r e s s .

TABLE 7 . 1

L IFE TABLE FOR THE TOTAL POPULATION

Age

d 'x

0 -1 1 -2 2 -3 3 -4 4 -5 5 -10 1 0 -15 1 5 -20 2 0 -25 2 5 -35 3 5 -45 4 5 -55 5 5 -65

2 6 1 4 1 9 2 1 1 9 13 6 5 5 8 4 2 2 33 2 04 1 43 2 1 014

d x

l x

q x

L x

2 . 6 1 . 4 1 . 8 2 . 1 1 . 9 1 .1 6 . 4 5 . 7 4 . 1 2 3 .0 2 0 .1 1 4 .1 5 . 7

1 00 9 7 .4 9 6 9 4 .2 9 2 .1 2 0 .2 7 9 .1 7 2 .7 6 7 .0 6 2 .9 3 9 .9 1 9 .8 5 . 7

. 0 26 . 0 14 . 0 18 . 0 22 . 0 21 . 1 23 . 0 81 . 0 78 . 0 61 . 3 66 . 5 04 . 7 12 1 . 0

9 8 .7 9 6 .7 9 5 .1 9 3 .2 9 1 .2 4 23 .3 3 79 .5 3 49 .5 3 24 .8 5 14 2 98 .5 1 27 .5 2 8 .5

7 5

T x 2 920 .5 2 82 1 .8 2 725 .1 2 630 2 536 .8 2 445 .6 2 022 .3 1 642 .8 1 293 .3 9 68 .5 4 54 .5 1 56 2 8 .5

2 9 .2 2 9 .0 2 8 .4 2 8 .0 2 7 .5 2 7 .1 2 5 .6 2 2 .6 1 9 .3 1 5 .4 1 .4 7 . 9 5

TABLE 7 . 2

ASSOCIAT ION BET WEEN BUR IAL AREA AND AGE AT DEATH FOR S UBADULTS

N o r th

0 1 1 20

A l lO the rA rea s

1 2

1 5

19

1 89

1 3 1

2 04

TOTAL

2 7 3 08

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w 0 0
50% o f r e po r ted c a se s ) , t h e s hou lde ra nd u ppe ra rm ,a nd t h ef o o t . A l lo ft h e se a r e s ugge s t ive o f a c t iv i ty i n duced d egene r a t ive d i sea se . F o r t h e s u cceed ing N eo l i th i c , t h e i n c idence o fa r th r i t i s i sr e duced , c ompa t ib le w i th w o r ldw ide f i gu re s f o r a r th r i t i s n o ted b y C oh en a nd A rme lago s ( 1 984a ) . T h i s s ugge s t s ad ec r ea se i n o ve r a l l w o rk load . R e la t ive ly l i t t le o t he r p o s t c r an ia l p a tho logy o ccu r s w i th in t h e Me so l i th ic . W i th t h e p o s s ib le e xcep t ion o f t h e I r on G a te s s i t e ( Ga r a shan in , 1 9 78 ; 2 i n a n o v i c , 1 9 75 ) , t h e l i m i ted e v idence f o ri n f e c t iou s d i sea se s ugge s t s al a ck o ft h ei n f ec t iou s " r e ser vo i r " t h a ti s s o t y p ica l o f u r ban i sed c ond i t ion s ( s e e A nd e r son a nd May , 1 979 ; May a nd A nde r son , 1 9 79 f o r a g ene ra l ized d i scu s s ion o f t h e d ynam ic s o f i n f e c t iou sd i sea se ) . A t f i r s t , c r an ia l p a tho logy w ou ld a l so a pp ea r i n f requen t i n b o th Me so l i th i c a nd N eo l i th i c E u rop e . R ecen t e v idence , h oweve r , s u gge s t s t h e e me rgence o f ad ua l p a t te rn o fp a tho logy i nt h e Me so l i th ic . O ne a r ea o f c r an ia l p a tho logy w i th a ppa r en t h i ghe r i n c idence t h an h a s b een p r ev ious ly r e po r ted i s c r an ia l p o ro t i c h ype ro s to s i s . P r ev iou s w o rk p r e sen ted o n ly t wo e xamp le s ( Cuzou l d e G r ama t , F ran ce , a nd F a lken s te inhoh le , G e rmany [ Me ik le john e t a l , 1 984 ] ) . H oweve r , r e cen t w o rk w i th D an i sh s amp le s , p r ima r i ly ( t hough n o t e n t i r e ly ) i n E r teb0 1 1e c on tex t s , s ugge s t s t h a t i t i s n o t u ncommon ( e .g . B Ogebakken , B e rgman sda l , B lok sb je rg a nd L undby ) . T he p r e sen ce o fp o ro t ic h ype ro s to s i s h a s a l so b een n o t ed a t F r an ch th i C ave ( Ange l , 1 984 ) . Wh i le n o p o ro t ic h ype ro s to s i s h a s b een n o ted a t V la sa c ( [ I ron G a te s o f t h e D anube ] N eme ske r i a nd L engye l , 1 978 ) , A nge l ( 1 984 ) i mp l ie s t h a t i tw a s p r e sen t . P o ro t i c h ype ro s to s i s i s s e conda ry t o r e duced

1 30

o f t h e l i ve r t oc onve r t v i tam in D i n to a n a c t ive a g en t . T he i mmed ia te e f f ec t w ou ld b e t h e l o s s o f v i t am in D a nd c a lc ium f r om t h e i n te s t ine d u r ing n o rma le l im ina t ion . T he e nd r e su l t , t h e s e conda ry l o s s o f c a lc ium t h rough t h e k i dney a nd i t s a b s ence f o r r e so rp t ion i n t h e i n t e s t ine , w ou ld a gg rava te t h e g ene ra l c ond i t ion o f p a r a s i t i sm . I f t h e i mp l ica ted t a pewo rm i s D iphy l lobo thr ium , w h ich h a s t h e d og a s a n i mpo r tan t i n te rmed ia te h o s t , t h en a f u r the r e f f e c t i s V i tam in B 12 d e f i c iency , w h ich l e ad s t ol owe r ed h a emog lob in p r oduc t ion , a nd t h e r efo r e t oa naem ia ( M a rqu a rd t a nd D ema re e , 1 985 ; B og i t sh a nd C heng , 1 990 ) . D iphy l lobo thr ium i s af r e sh wa te rp a r a s i te ,f ound i n , a mong o t he r s , p i ke , s a lmon a nd t r ou t . S cand inav ia a nd t h e D anub eb a s in a r e b o th a r ea s w he r e t h i s p a r a s i te i se ndem ic . T hu s , s i gn i f ican t l e ve l s o f i n te s t ina l p a ra s i te s , a mong f i sh ing c ommun i t ie so fn o r ther na nd e a s te rn E u rope ,m ay h ave c omp rom i sed a no t he rw i se h i gh q ua l i ty d i e t , i nt e rms o fb ene f i tt ot h ei n d iv idua l . We wou ld a l so l i ke t o p o in t o u t af u r the r p o s s ib le me chan i sm f o r t h e a pp ea rance o f b one s t r e s s ma rke r s i n t h e p r e sence o f ah i gh m a r ine d i e t . L azenby a nd McCo rmack ( 1 985 ) p o in t o u t t h a t h i gh u sage o f s a lmon c an r e su l t i n h ype rv i tam ino s i s -D . S i nce v i tam in D c an s t imu la te c a lc ium w i thd rawa l f r om t h e s k e le ton , t h i s p rov ide s ab a s i s f o r m a rke r s s u ch a sp o ro t ic h ype ro s to s i s . I ts hou ld a l so b e p o in t ed o u t t h a t h ype rv i tam ino s i s i s ap o ten t ia l ly f a ta l s ynd rome , a nd t h a t t h e s i tua t ion o u t l ined b y L azenby a nd M cCo rmack i st h u s a n e x ten s ion o f t h e p o ten t ia l f o r p ro te in p o i son ing t h a t c an r e su l t f r om e x ce s s ive u s e o f m a r ine r e sou r ce s ( No l i a nd A ve ry , 1 988 ) . As im i la rp a t te rn o fi n su l t sm ay h ave o ccu r red a s ar e su l t o f h i ghe r p opu la t ion d en s i ty a nd i n c r ea sed s e den t i sm o f t h e se p opu la t ion s , i f h ype ro s to s i si st a ken a sa ni n d i ca to ro fh i ghe rr a te o f i n f ec t ion ( i . e . K en t , 1 9 86 ) . T he r e i s s t rong c u l tu ra l e v id ence , i nt e rm s o fs e a sona l i n d i ca to r s o f o ccupa t ion a nd t h e p e rmanen t n a tu r e o f d we l l ing s , t h a t b o th t h e i n hab i tan t s o f t h e I r on G a te s a nd t h e p opu la t ion s o f t h e E r t ebo l le w e re m o r e s eden ta ry t h an o t h e r Me so l i th i c g roup s ( Row ley -Conwy , 1 983 ; P r i ce , 1 985 ; P r in tz , 1 987 ; C la son , 1 980 ; B o lom ley , 1 973 ; V oy t ek a nd T r ingham , 1 989 ; C hapma n , 1 989 , e t c ) , a l though w er e cogn i ze , i nt h i s c on tex t , t h a t s e den t i sm d oe s n o t n ece s sa r i ly m ean y e a r round p e rmanen t h ab i ta t ion o f as i ng le s i t e . S ub -op t ima l l e ve l s o f h ea l th c an b e a l so d e t ec ted t h rough t h e p r e s ence o f d en ta l e name l h ypop la s ia . P rev iou s w o rk h ad s ugge s ted t h a t t h e t r a i t w a s a lmo s t u n ive r sa l ly a b sen t i n t h e E u ropean Me so l i th ic a nd N eo l i th ic ( Me ik le john e ta l , 1 984 ) . Mo r e r e cen t w o rk o n b o th D an i sh a nd P o r tuge se c o l lec t ion si n d i ca te st h a tl ow l e ve l s o f e name l h ypop la s ia a r e p r e sen t i n b o th c a se s ( A lexande r sen , 1 988 ; L ub e l l a nd J a cke s , 1 989 ; L ub e l l e ta l , 1 989 ) , a l though o n ly af ew c a se sa r e o ft h es eve r et y pe r epo r ted i ns ome a r ea so fN o r th

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Ame r ica , f o r e xamp le . I n c on t ra s t , y 'Edynak ( 1 989 ) r e po r t s t h ep r e sence o fe name l h ypop la s ia o n7 0% o fa l lt e e th a tV la sa c . T he l e ve l sr e co rded f o rt h es e cond mo la r sa r et a ken a sa n i n d ica t ion o f t h e s e ve r i ty o f t h e c ond i t ion . H ypop la s ia i sa l so p r e sen t a tP ad ina ( y 'Edynak , 1 978 ) . I n g ene r a l , s t ud ie s o f e name l h ypop la s ia l e ve l s a c ro s s t h e Me so l i th i c -Neo l i th ic t r an s i t ion h ave n o t b e en c a r r ied o u t . O ne c a se , h oweve r , i s t h a t o f A lexande r sen f o r s ou the rn S cand ianav ia ( 1 988 ) . I n t h i s c a se h e h a s s hown t h a t 5 3 .5% o f t h e i n d iv idua l s f r om V edbaek -B0gebakk en , S ka teho lm I a nd S ka teho lm I s how m i ld h ypop la s ia ( n=58 ) . I nc ompa r i son ,4 5% o fD an i sh m idd le a nd l a te N eo l i th i c i n d iv idua l s s how t h e t r a i t ( n=102 ) . T he d i f f e rence i s n o t s i gn i f ican t ( c h i squa r e= 1 .03 , p = .3095 ) . I ts hou ld b e s t re ssed , h oweve r , t h a t t h e d i e ta ry b a se o f t h e se t wo s amp le si sp ro found ly d i f f e ren t ,a ss hown b yb one i s o tope s t ud ie s ( Taub e r , 1 981 ; P r ice , 1 989 ) . F o r t h i sr e a son w ec anno ta s sume t h a tt h ec au se o ft h e h ypop la s ia i s i d en t ica l i n t h e t wo c a se s . U n fo r tuna t e ly , as im i la r e nume ra t ion o f e name l h ypop la s ia f o r o t he r r e g ion s i nt h e N eo l i th ic h a s n o t y e t b een c a r r ied o u t , m ak ing i ti mpo s s ib le a t t h i s t i me t o m ak e g ene ra l iza t ion s a bou t t h e i n c idence o f h ypop la s ia a c ro s s t h e a g r icu l tu ra l t r an s i t ion . T he a bove d i s cu s s ion s ugge s t s ad ua l p a t te rn i nt h e p a t te rn s o ft h e se p a tho log ica l c ond i t ion s i n E u rop e : b o th p o ro t ic h ype ro s to s i s a nd e name l h ypop la s ia a r e mo r e c ommon t h an h a s b een b e l ieved , b u t o ccu r p r inc ipa l ly o n s e t t lemen t s i n e a s te rn a nd n o r the rn E u rope . H e r e t h ey a r e a s so c ia t ed s t rong ly w i th f i sh ing a nd i n c re sed r e s iden t ia l p e rmanen ce , t h u s l e ad ing u s t o a s e cond p r opo s i t ion : P ropo s i t ion 2 : T he r e i se v idence f o r s pa t ia l v a r ia t ion a nd m a jo r d i e ta ry d i f f e r en ce s i n l a te Me so l i th i cp opu la t ion sp r io rt ot h ei n t roduc t ion o f f a rm ing . T h i s p ropo s i t ion c an b e b e s t i l u s t ra ted b y d en ta l p a tho logy . T he mo s t w ide ly r e po r ted d en ta lp a tho logy i nE u rop eh a sb een c a r ie s . I nt h e e a r l ie rs u r vey ( Me ik le john e ta l , 1 984 ) Me so l i th ic c a r ie s l e ve l s w e r e r epo r ted a s l ow c ompa red t o N eo l i th i c l e ve l s ( 1 .9% v s 4 . 2% ) . H oweve r , mo re r e cen t w o rk s ugge s t s am o r e c omp lex p a t ter n , i n c lud ing a l a ck o f g eog r aph ic u n ifo rm i ty ( Me ik le john e ta l , 1 988 ; F raye r , 1 989 ) . C u r ren t o ve ra l lr e su l t sa r ep r e sen ted i nT ab le 1 .T he e a r l ie r r e po r ted f i gu r e s c l ea r ly u nde r r epo r ted c a r ie sr a t e s f o r t h e Med i te r ranean r e g ion . T he o ve ra l l p a t ter n i sc l ea r , w i th a ve rage c a r ie s r a te s i n c r ea s ing f r om t h e B a l t ic t h rough m id la t i tude E u rope t o t h e Med i te r ranean b a s in .T he e x cep t ion t ot h i sp a t t e rn i so n ce a ga in a tV la sac a nd P ad ina i nt h e D anub e G o rge s , w he re a b sence o f c a r ie s ( F r aye r , 1 989 ; inanov ic , 1 975 ) a l ign s t h i s s e t t lemen t w i th t h e B a l t ic s i t e s r a the r t h an t om id la t i tude s i te s s u ch a sT ev ie c a nd H oed ic . T he d i f f e r ence s a r e h i gh ly s i gn if i can t . Wha td ot h ey s ugge s t ?

C r i t ica l t oc a r ie s d eve lopmen t i sc a rbohyd ra te c on sump t ion . D ie ta ry c a rbohyd ra te a f f ec t sp l aque p H , i n c rea s ing a c id i ty t ot h e p o in t w he r e d en ta l e name l i s d i s so lved . D ie ta ry p ro te in r e su l t s i n i n c r ea sed a l ka l in i ty o f t h e p l aque , l e ad ing t o m ine ra l c r y s ta l isa t ion a nd c a l cu lu s . T he p r e sence o fc a lcu lu sa nd c a r ie si n d ica te a l t e rna te p e r iod so f a c id i ty a nd a l ka l in i ty a nd t h e a l te rna t ion o f c a rbohyd r a te a nd p ro te in d i e ta ry s ou r ce s . ( Fo r m o r e c omp rehen s ive r e v iew s o fc a r ie s f o rma t ion , s e e H i l l son ( 1 986 ) , Me ik le john e ta l( 1 988 ) a nd F raye r( 1 989 )a nd t h e i rc on ta ined s ou r ce s ) . F rom t h e a bove , i tc an b e s e en t h a t t h e i n c idence o fc a r ie s i si ni n ve r se r e la t ion sh ip t ot h e b imoda l p a t ter n o f o t he r p a tho log ie s , n o t ed a bove . T he d a ta f r om V la sa c , P ad ina a nd t h e S cand inav ian m a te r ia l s s how c a lcu lu s , b u t c a r ie s a r e v i r tua l ly a b sen t . A p ro t e in -ba sed d i e t i s s uppo r ted , a c onc lu s ion b o l s te r ed b y f a una l a na ly s i s f o r t h e t wo a r ea s , a nd b one c hem i ca l a na ly s i s f o r t h e B a l t ic s i te s . A s s hown a bove , t h e se s i te s a l so h ave ah i ghe r i n c idence o fo t he r p a tho logy . I n t h e Med i te r ranean s amp le s , b o th c a lcu lu s a nd c a r ie sa r e p r e sen t , s ugge s t ing am ixed p ro t e in a nd c a rbohyd ra te d i e t . A ga in , t h i s i ss uppo r ted b y p a la eoeconom ic a nd b one c hem i ca l a na ly s i s a t s i t e s s u ch a sU zzo , w he re t r ace e l emen t a na ly se s g ene ra l ly s ugge s t a c a rn ivo rou s d i e t a nd p a r t icu la r ly h i gh c on sump t ion o f m o l lu s cs , w h i le r e la t ive ly h i gh i n c idence o f c a r ie s a nd t h e p r e sen ce o f m a c robo tan ica l r ema in s i n d ica t e a d i e t e ry c omponen t o f f r u i t s , n u t s a nd b e r r ie s ( Me ik le john e ta l , 1 988 ; B o rgogn in i -Ta r l i , 1 9 89 ; F raye r , 1 989 ) . Ma c ro scop ic f i nd s o f l e gume s , n u t s a nd f r u i t s w e r e a l so m ad e a t B a lma A beu r ado r i n F rance ( Vague r e t a l , 1 9 86 ) , F on tb regoua ( Cou r t in , 1 974 ) a nd e l sewhe r e i n t h e We s te rn Med i te r ranean ( B inde r , 1 989 )a sw e l l a s a t F ranch th i i n G r ee ce ( J a cob sen , 1 9 76 ; H an sen a nd R enf r ew , 1 9 78 ) . A t C ing le V e rme i l , i nC a ta lon ia , w he r e e i gh t o u t o f n i ne t e e th w e r e f o und t ob e c a r iou s , t h e a s so c ia ted c u l tu ra l l a ye r c on ta ined t h e r ema ins o fw a lnu t , p l ums , c he r r ie s , p i ne n u t sa nd a co rn s( Tu rbon , 1 989 ) .T he r ei sa l so e v id ence f o r t h e u s e o f h a ze l n u t s , c e r ea l s a nd C henopod ia a tV la sa c ( Ca r cu ima ru , 1 978 ; P r in tz , 1 987 ) , t h ough w i thou t c a r ie s , a nd a t I c oana ( Ca r cu ima ru , 1 978 ; V oy tek a nd T r ingham , 1 989 ) . T he s ugge s t ion o f a n i n c r ea sed c a rbohyd r a t e i n take i sf u r the r s uppo r ted b y d e l ta 1 3C f i gu r e s f r om P o r tuga l ( Mo i ta d oS eb e s t iao , 1 6 .4 ; C abe co d a A r ruda , 1 7 .6 ; N eo l i th ic - 1 9 .8 ) (Lube l l a nd J a cke s , 1 989 ) . T he i n c idence o f c a r ie s i n t h e We s t Med i t e r r anean L a te Me so l i th ic -9 . 45% -i sa bou t 4t i me s h i ghe r t h an t h e c a r ie s r a te a mong t h e N a tuf ian s o f 2 3% ( Sm i th e t a l , 1 984 ; S m i th , 1 989 ) a nd a bou t t w i ce a sh i gh a st h e c ompa r ab le f i gu re f o r r e po r ted l e ve l s f r om t h e E u rop ean N eo l i th i c . I ti sa l so h i ghe rt h an N eo l i th ic l e ve l si n t h eL evan t ( F raye r , 1 989 ; Me ik le john e ta l , 1 988 ; Sm i th e ta l , 1 984 ) a nd c an b e c ompa r ed t ot h e

l e ve l s a t t a ined i n t h e c a rbohyd r a t e o r ien t ed a g r icu l tu ra l c ommun i t ie s i n N o r th Ame r ica d u r ing t h e M i ss i s s ip ian a nd Wood land p e r iod s ( Cook , 1 984 ; P e r z ig ian e ta l , 1 984 ; R o se e ta l , 1 984 ) . C omp l emen ta ry d a ta c ome f r om a n e xam ina t ion o f t o o th s i ze i n t h e P o r tuge se Me so l i th ic s i t e s o f Mo i ta d o S eba s t iao a nd C abe co d aA r r uda . Ag ene ra ld ec rea s ei ns i ze w a s n o ted b e twe en t h e s i te s , ap e r iod o fo n ly h a lf a m i l lenn ium .T h i s t r end s e ems t oc on t inu e i n to t h e N eo l i th i c ( M e ik le john a nd S chen tag , 1 988 ) . H oweve r , t h e se c hange s b egan w i th in t h e Me so l i th ic a nd a r e t h e r e fo re n o t t h e p r oduc t o f t h ea g r icu l tu ra l t r an s i t ion p e rs e . As im i la rp a t te rn h a s b een o b se r ved a tt h e Y ugo s lav ian I r on G a te s i t e s , a ga in w i th in t h e Me so l i th ic ( yEdynak , 1 989 ) . I th a sb e en s ugge s ted t h a t t h e r e duc t ion i n p o s t e r io r t o o th s i ze , a nd i n t h e d en t i t ion i n g ene ra l , r ef l ec t s as h i f t t o a mo r e c a rbohyd ra te d i e t , u sua l ly v i ewed a st h e p rodu c t o fa g r icu l tu r e ( Ca lcagno a nd G ib son , 1 988 ) . H oweve r , a s i gn if ican t s h i f t t oa ny s o f t e rd i e t c ou ld l e ad t oa d e c r ea se i n m and ibu la r s i ze a nd i n d en t i t ion ( y 'Edyna l c , 1 9 78 , 1 989 ) . B ea r ing i n m ind a s so c ia ted e v id ence ,d en ta lr e duc t ion s u ppo r t s t h e c a se f o r as h i f tf r om m ea t -ba sed t oe i the rt h e f i sh o r ien ted o r c a rbohyd r a t e -o r ien ted d i e t i nt h e l a te M e so l i th ic . T hu s ad i e ta ry s h if t c an b e p o s i ted f r om e v iden ce f r om b o th d en ta l c a r ie s a nd d en ta l s i ze ( t hough F r aye r ( 1 989 ) a r gue s a g a in s t a n o ve r a l l t empo ra l t r end i nc a r ie s r a te s w i th in t h e Me so l i th ic ) . B a sed o n t h e se c on s ide ra t ions , ad ua l p a t te rn o fd i e t a nd l i f e s ty le s e em s t ob e e me rg ing f o r t h e E u ropean M e so l i th ic ( F ig . 1 ) . I n t h e w e s t Med i te r ra nean , ac a se c an b e m ad e f o r t h e c on sump t ion o f ab a lan ced d i e t , b a sed o n p l an t f o od s , f i sh a nd m ea t . T he g r ea te r i mpo r ta nce o fa c a rbohyd r a te d i e t i n t h e Me so l i th i c g i ve s s ome s uppo r t f o r t h e e x is tence o f i n d igenou s p l an t h u sb and ry , o r a t l e a s t as h if t t owa rd s mo r e i n ten s ive g a the r ing o f w i ld p l an t f o od s ( t hough t h e e v id ence f r om P o r tuga l i s f o r t h e l a te r Me so l i th ic , t h e d a ta f r om U zzo a nd Mo la ra i n S i c i ly d a te t ob e fo r e 7 000 b c ( Bo rgogn in i -Ta r l i e t a l , 1 989 ; C on s tan t in i , 1 989 a nd c on ta ined r e f e r ence s ) . S uch ap a t te rn h a s b een d e s c r ibed b y C ohen ( 1 977 , 1 989 ) a nd B in fo rd ( 1968 , 1 9 83 ) a s as h i f t f r om p r e f e r r ed t o s ub -op t ima l r e sou r ce s , a nd t h e r e fo r e a s a n i n d i ca t ion o f p opu la t ion p r e s su re a nd d emog raph ic s t re s s . A nd y e t , w i th t h ee xcep t ion o fc a r ie s , t h e i n d ica t ion so fp hy s ica l s t r e s s a r e l owe r i n t h i s p a r t o f E u rop e t h an e l sewhe re . I n t h e s o u th B a l t ic a nd a t L ep en sk i V i r , a no the r d i e te r y p a t te rn i s e me rg ing ( F ig . 1 ) , w h i ch i s b a s ed o n t h e g r ea t s i gn i f i can ce o f a qua t ic r e sou r ce s . F auna l a nd t e chno log i ca l e v idence f r om o t he r s i te s i nn o r thern a nd e a s te rn E u rope s ugge s t t h a t t h i s h a s b een t h e p r eva len t p a t t e rn o ve rm uch o ft h is a r ea ( C la rk , 1 9 76 , 1 952 ; L oze , 1 979 ; T r ingham , 1 973 ; Z ve leb i l , 1 9 86a ,b ;

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p opu la t ion ( s ee G oodman e t a / , 1 984b , t h i s v o lume ) . S t a tu r e e s t ima t ion s f o r E u ropean s amp le s f r om t h e U ppe r P a laeo l i th ic t o t h e N eo l i th i c h ave b e en r e po r ted b y Me ik le john e ta l ( 1 984 ) . F rom t h e U ppe r P a laeo l i th ic u n t i l t h e N eo l i th ic , p opu la t ion s b ecome p rog r e s s ive ly s ho r te r . O n ly a f te r t h e l a te N eo l i th ic i st h e t r end r e ve r sed . T he d i f f e r ence b e tween t h e a ve rage Me so l i th i c a nd a ve rage N eo l i th ic s t a tu re d oe s n o t r ef l ec t a b r up t c hange . I n f a c t , o ve ra l l Me so l i th ic ( n.82 ) a nd N eo l i th ic ( n= 190 ) s amp le m ean s a r e s t r ik ing ly s im i la r ; 1 67 .7 a nd 1 67 .3 c m f o r m a le s , 1 55 .6 a nd 1 54 .1 c m f o r f ema le s . Wha teve r i s p roduc ing s ho r ts t a tu r e i nt h e Me so l i th ic i ss t i l la f a c to r i n t h e N eo l i th i c . I f r e duc t ion i n s t a tu re i n d ica te s n u t r i t iona l s t r e s s , a ss ome h ave c l a imed , t h en t h e s t re s s w a s n o t r e so lved u n t i l t h e e nd o f t h e N eo l i th i c . T h i s w ou ld s uppo r t e v idence t h a t , w h i le t h e a dop t ion / sp read o f f a rm ing e conomy m ay b e , i np a r t , ar e spon se t op r e s su re s s e t u p i n l a te Me so l i th ic t i me s , i td oe s n o t f u l ly a l lev ia te t h em ( s ee C ohen a nd A rme lago s , 1 984a , R oo seve l t , 1 984 ) . F a rm ing m ay s imp ly a low m o r e p eop le t o s u rv ive w i thou t i mp rov ing t h e i r g ene ra lh ea l th .

R ow ley -Conwy a nd Z ve leb i l , 1 989 ) . H uman s k e le ta l a na ly s i s s ugge s t s t h a t t h i s " no r th -ea s t " E u rop ean p a t te rn w a s a s so c ia ted w i th g r ea t e r l e ve l s o f p a tho logy a nd p hy s ica l s t r e s s t h an w a s t h e c a se w i th t h e "We s t Med i t e r r anean " p a t t e rn . T he p a tho logy m ay b e r ef lec t ive o f i n c r ea sed s e a son a l v a r ia t ion i n t h e f o od s upp ly , i n c r ea sed r e s id en t ia l p e rmanence , a nd /o r p r e sence o f p a r a s i t ic d i sea se a r i s ing f r om c on sump t ion o f c on tam ina ted f i sh . P ropo s i t ion 3 : We s ugge s tt h a to n t h ep r e sen t e v iden ce , l i t t le o r n o d i f f e rence e x i s t s i n t h e o v e r a l l h ea l th s t a tu s o f Me so l i th i c a nd N eo l i th ic p opu l a t ions . A dm i t ted ly , t h i s i s ad i f f icu l t p ropo s i t ion t o e va lua te a tp r e sen t ,b e cau se o fw ide d i s c r epan c ie s i nt h e a pp roa ch , t h e l e ve l o fa na ly s i s a nd t h e s i ze a nd t h e d i s t r ibu t ion o f s k e le t a l s amp le s r e p r e s en t ing Me so l i th ic a nd N eo l i th ic p opu la t ions . A t p r e sen t , t h e re h a s b een m o r e e x ten s ive a na ly s i s o f Me so l i th i c t h an o fN eo l i th ic r ema in s i n am anne r a low ing f o r a n a c cu ra te r e con s t ruc t ion o f h ea l th s t a tu s . Me ik le john e ta l ( 1 984 ) h ave d i s cu s sed t h e p r ob lem o f d e r iv ing s u ch i n fo rma t ion f r om t h e N eo l i th ic d a ta a va i lab le . T h i s a ppea r s t ob e ar ef l ec t ion o f t h e c l in i ca l a pp ro ach t o t h e d e s c r ip t ion o f h uman r ema in s ( a s d i scu s sed a bove ) . I ta l so i s ,n od oub t , ar ef l e c t ion o f t h e t r ad i t iona l v i ew , w he re s t r e s s w a sn o te xpe c ted a mong f a rme r s ,w he r ea s ,a mong f o r age r s , d i e ta ry s t r e s s w a sa n t ic ipa ted . T h i s v i ew f i t s w i th e i the r o f t wo m od e l s , t h e c l a s s i c " poo r s t a rv ing s avage s " c a r ica tu r e o fp r e -Neo l i th ic l i f e , o r w i th s t re s s a s t h e " p r ime m ove r " o f t h e t r an s i t ion t o f a rm ing a s p e r C ohen ( 1 977 ) . T he e nd r e su l t i s t h a t w e c an o n ly s umma r i se t h e c u r r en t s t a te o f t h e e v idence , t h e c ompa r a t ive v a lue o fw h ich i sl i m i ted . F o r e xamp le , t h e p r e s en t e v idence s how s h i ghe r l e ve l s o f a r th r i t i s , e name l h ypop la s ia a nd p o ro t ic h ype ro s to s i s i n t h e Me so l i th i c , a l though t h e l a t te r t wo m igh t h ave b een u nde r - r e po r t ed i n t h e N eo l i th ic s amp le s ( Me ik le john e ta l , 1 984 ) . YEdynak ( 1978 ) n o t e s t h a te name l h ypop la s ia a t P ad ina ( I ron G a te s ) w a s mo r e c ommon i n M e so l i th ic t h an i n N eo l i th i c l e ve l s . P ap ( 1 989 ) r e po r t st h e p r e sence o fp o ro t ic h ype ro s to s i s i nt h e e a r ly a nd l a te N eo l i th ic o fH unga ry , w h i le n o t ing t h e a b sence o f e name l h ypop la s ia . Many o t he r r e po r t s d o n o t m en t ion e i the r c ond i t ion ( Me ik le john e ta l , 1 984 ) . S im i la r ly , i t w a s t h ough t t h a t c a r ie s f r equenc ie s w e re h i ghe r i nt h e N eo l i th ic t h a n t h e M e so l i th ic ( s e e a bove ) . H oweve r , t h e s i tua t ion i s f a r m o re c omp lex . A s as imp le e xamp le , t h e r e ca l cu la ted o ve ra l l Me so l i th ic c a r ie s r a te f r om T ab le 1 .1 i s5 . 03% , h i ghe r t h an t h e p r ev iou s ly n o ted f i gu re o f4 . 2% , d e r ived f r om 1 6 N eo l i th ic s i t e s f r om t h e N e the r land s , B e lg ium , F ran ce a nd P o r tuga l ( s ee a bove , a nd a ppend ix t o Me ik le john e ta l , 1 984 ) . S t a tu r e i sc l e a r ly a m a rke r f o r t h e o ve r a l l h ea l th a nd n u t r i t iona l s t a tu s o f a

P ropo s i t ion 4 : t h e r e i se v idence f o r i n cr ea se i n p opu la t ion d en s i ty i n s ome a r ea s o f l a t e Me so l i th i c E u rope , p a r t i cu la r ly i n N o r the rn a nd E a s t e rn E u rope . I na r ea s s u ch a s s ou the rn S cand inav ia o r t h e n o r th -ea s t B a l t i c , a nd i n t h e N o r th E u ropean P l a in ,t h ec u l tu ra le v idence c ons i s t so fa n i n c r ea se i n t h e s i ze a nd p e rmanence o f s e t t lemen t s , d ec r ea se i nt h e s i ze o fa ppa r en t t e r r i to r ie s a nd t h e a ppea ra nce o f c eme ta r ie s ( F ig . 1 ) ( Br inch P e te r sen , 1 973 ; C hamp ion e ta l , 1 984 ; N ewe l l , 1 984 ; C on s tand se - We s t e rmann e t a l , 1 984 ; Z ve leb i l a nd R ow ley -Conwy , 1 986 ) . T he se c u l tu ra l d eve lopmen t s a pp ea r t o b e r ef le c ted i n t h e b i o log ica l r e co rd , a l though t h e e v idence r ema in s t e nuou s . T he t e r r i to r ia l b ehav iou r a nd d en s i ty i n c r ea se a r e m i r ro red i n t h e n umbe r o f s k e le ton s s how ing s i gn s o f v i o lence . T ype s o f i n ju ry i n c lude a ppa r en t b l ow s t o t h e s ku l l , p o s s ib ly c au sed b y h eavy s t r ik ing w eapon s , a nd e mbedd ed p ro jec t i le p o in t s ,m o s t f r equen t ly i nt h e v e r t eb ra l c o lumn ( B r inch P e te r sen , 1 981 ; N ewe l l , 1 984 ;C on s tand se - We s te rmann a nd N ewe l l , 1 984 ; Me ik le john e t a l , 1 984 ; A nge l , 1 984 ; P r in tz , 1 987 ) . I fs u ch b ehav iou ri sd en s i ty d ependan t , a s h a s b een s ugge s ted b y w o rke r s s u ch a s C hagnon ( 1972 ) , s e e a l so H yman a nd S cu t t , t h i s v o lume ) , t h en w e h ave a b i o log i ca l m a rke r f o r t h i s d emog raph ic s h i f t . A s n o ted a bove , d en s i ty i n c r ea se m ay a l so b ei mp l ica ted i nt h ep r e sence o f c ond i t ion s s u ch a s p o ro t ic h ype ro s to s i s a nd , p o s s ib ly ,e name lh ypop la s ia . A no the rt r end s ugge s t ive o f i n c rea sed d en s i ty d u r ing t h e l a t e Me so l i th ic i s t h e i n c rea sed v a r iab i l i ty a nd g r ea te rc i r cum scr ip t ion i np hy s i ca l c ha ra c te r i s t i cs i n t h e se p opu la t ions . T h i s m ay s ugge s t t h a t as h i f t h a s t a ken p l a ce f r om " b and

1 3 3

l e ve l " s o c ie t ie s w i th o p en m a t ing n e two rk s ( Me ik le john , 1 974 , 1 977 ) t o mo r e c i r cum s c r ibed s o c ie t ie s w i th g r ea te r g ene t ic i s o la t ion ( Newe l l , 1 984 ; C on s tand se - We s t e rmann e t a l , 1 984 ) . F ina l ly , a n a na ly s i s o f t h e d i s t r ibu t ion o f o r namen t s i n E u ropean s i te s s ugge s t s a n i n c r ea sed c i r cums c r ip t ion a nd i n teg ra t ion o f c u l tu ra l u n i t s o ve r t h e d u ra t ion o ft h e Me so l i th i c , o ccu r r ing e a r l ie s t i n t h e s ou th a nd s ou th -ea s t ( Con s tand se - We s t e rmann & N ewe l l , 1 988 ) . H oweve ri n t e rp r e ted , t h i se v idence a tt h e v e ry l e a s t p o in t s t or e g iona l v a r iab i l i ty i n p opu la t ion n umbe r s d u r ing t h e l a t e Me so l i th ic a nd i mp l ie s t h a ta tl e a s ti ns ome a r ea s ,p opu la t ion d en s i ty m ay h ave b een a s h i gh a s i tw a s i n t h e s u cceed ing e a r ly N eo l i th ic .

N e e ley , 1 9 87 ; C on s tand se -W e s t e rmann a nd N ewe l l , 1 989 ) , o f a s c r ibed s t a tu s ( O 'Shea a nd Z ve leb i l , 1 9 84 ; C la rk a nd N ee ley , 1 987 ) a nd o f s pe c ia l , o c cupa t iona l ly re la ted s t a tu se s ( O 'Shea a nd Z ve leb i l , 1 984 ) h a s b een n o t ed . I ng ene r a l i t s e em s t h a t b o th m en a nd w omen c ou ld a cqu i r e h i gh r a nk i n Me so l i th ic s o c ie ty ( O 'Shea a nd Z ve leb i l , 1 984 ; C on s tand se - We s t e rmann a nd N ewe l l , 1 989 ) a l though m en t e nded t o a qu i r e h i ghe r r a nk mo r e o f ten t h en w omen ( O 'Shea a nd Z ve leb i l , 1 9 84 ) . C on s tand se - We s t e rman a nd N ewe l l ( 1 989 ) e xam ined s ke le ta l m a rke r s o f m echan ica l s t r e s s i n t h e l i gh t o f t h e s o c ia l d i f f e r ence s n o ted a bove a nd f o und t h a t h i ghe r r a nk ing m a l e s s how s t ronge r l i mb l a te ra l i sa t ion t h an t h o se i nl owe r r a nk s , w h i l e h i ghe r r a nk ing f ema le s s how l e s s l i mb l a te ra l isa t ion a nd a r th r i t i s t h an l owe r r a nk s . T he m ean ing o f t h e s e d i f f e r ence s r ema in s u nc lea r ; o ne p o s s ib le e xp lana t ion m ay b e t h a t m a le s a cqu i r ed s t a tu s t h rough s p e c ia l i sed p hy s i ca l a c t iv i ty , w h i le h i gh f ema le s t a tu s w a s a s c r ibed b y d e scen t , b y a s so c ia t ion w i th d om inan t ma le s , o r t h rough a c t iv i ty w h i ch d i d n o t l e ave p hy s ica l m a rke r s o n t h es k e l e ton . B ea r ing i nm ind t h e i mponde r ab le s o c ia l a nd i d eo log i ca l c ode s w h ich g overn t h e r e la t ion sh ip s b e twe en t h e l i v ing a nd t h e d ead , a nd t h e b u r ia l a nd e vo ca t ion o f t h e l a t t e r , much o f t h e a bov e d i s cu s s ion i s , p e r f o r ce , s pe cu la t ive . T he r e i s e nough t e n ta t ive e v idence a tt h i s s t age , h oweve r , t os how t h a ts o c ia l d i f f e r en t ia t ion d i d e x i s t i nt h e l a t e Me so l i th i c i nm o s t p a r t s o f E u rop e , a nd t h a t t h i s m ay h av e s e rved a sa n a dd i t iona l v a r iab le i n t h e d e te rm ina t ion o fh ea l th a nd p a tho logy a mong t h e se p opu la t ion s .

P ropo s i t ion 5 : T he r e i s e v idence f o r d i f f e r en ce s i nh ea l th a nd s o c ia l s t a tu s a mong t h e i n hab i tan t so f Me so l i th ic s e t t lemen t s . T he h uman b i o log i ca l e v idence p o in t s t o d i f f e r ence s i nh ea l th s t a tu s a nd p a t te rn o f i n su l t s b e twe en m a le s a nd f ema le s , w h ich , a l though c on t ingen t t o s ome e x ten t o n t h e s e x o f i n d iv idua l s a nd t h e c h i ldbea r ing r o le o f w omen ( s ee B u sh ,t h i sv o lume ) ,m ay a dd i t iona l ly i n d i ca te a n u nequa l a c ce s s t o r e sou r ce s , w h ich , o n t h e w ho le , p r e jud iced t h e h ea l th s t a tu s o f f ema le s . F ema le s a ppea r t o s u f f e r f r om g r ea te r i n c idence o f d en ta l d i sea se a nd n u t r ien t d e f ic iency . F raye r ( 1 989 :262 ) n o t ed t h e g r ea te r i n c idence o f c a r ie s a mong f ema le s : " F ema le c a r ie s r a te s a r e a bou t d oub le t h o se f o r t h e m a le s a nd s i gn i f ican t ly d i f f e ren ta tt h e0 . 007 l e ve lo rb e t te r " . A t V la sac , f ema le s s u f f e red f r om m o re s e ve r e l e ve l s o f a l veo la r r e so rp t ion a nd o s teoma la c ia , t h ough m a le s h ad ah i ghe r i n c idence o fc a lcu lu s ( y 'Edynak , 1 989 ) . B o th o ft h e se d a ta s e t s i n d ica t e ag r ea te r l a ck o f c a lc ium , p r o te in a nd p o s s ib ly o t he r n u t r ien t s a mong f ema le s , w h ich c ou ld b e c au sed , a t l e a s t i n p a r t , b y p r egnancy a nd l a c ta t ion . T he q ue s t ion i s w he the r t h e se d i f f e r ence s c an b e e xp la ined b y c h i ldbea r ing a l one . T he d i f f e r en t ia l p a t te rn i nt h ei n c idence o f c a r ie s a nd c a lcu lu s s ugge s t s t h a t ad i e ta ry f a c to r m ay h ave b een a dd i t iona l ly i mp l ica ted : m a le s c on sumed m o r e p ro t e in a nd c a lc ium , w h i le f ema le s c oncen t r a ted m o re o n ac a rbohy ra te d i e t ( y 'Edynak , 1 989 ) .T h i sp a t ter nf i nd ss ome s uppo r t i n w ha t w e k now o f f o od s ha r ing p r a c t ice s a nd f o od p rocu remen t a mong mode rn h un t ing a nd g a the r ing p eop le , w he r e m a le s m ay b e a b le t o c ap i ta l ize o n t h e c on sump t ion o f t h e m o s t n u t r i t iou s m ea t p a r t s ( Spe th , 1 989 , 1 990 ; K en t , 1 989 ) . Ac omp r ehen s ive t r a ce e l emen ta na ly s i so f s ke le ta ls amp le ss hou ld r e so lve t h i sq ue s t ion . C u l tu ra l e v idence f o r t h e e x i s tence o f s o c ia l r a nk ing i sb a sed m a in ly o n t h e e xam ina t ion o f m o r tua ry c on ten t s f r om Me so l i th ic c eme ta r ie s . T he e x i s tence o f g ene ra l i sed w ea l th o r s t a tu s r a nk s ( O 'Shea a nd Z ve leb i l , 1 984 ; C la rk a nd

P ropo s i t ion 6 : N e i the r s ke le ta l m o rpho logy , n o r g ene t ic i n fo rma t ion d e r ived f r om c u r r en t E u ropean p opu la t ion s g i ve s c l ea r s uppo r t t o t h e c o lon i sa t ion m od e lf o rt h et r ans i t ion t of a rm ing . We w ou ld a r gue t h a t t h e s ke le ta l m o rpho logy o fN eo l i th i c p opu la t ions s how s ac on t inua t ion o f t r end s t h a tw e re a l r eady p r e sen ti nt h e Me so l i th ic . T h i s i st h e c a se w i th c hange s i n d en ta l s i ze a nd a nd i nt h e r e duc t ion i ns t a tu re , r e f e r r ed t oa bove . T he s i gn i f i ca nce o ft h i s o b serva t ion i st h a t t r end s t h a t a r e a l ready v i s ib le i nt h e Me so l i th ic c anno t b e u s ed a s e v idence f o r t h e e f f ec t s o f t h e N eo l i th i c " r evo lu t ion " . A s ar e g iona l s amp le , c r an ia l m a t e r ia l s f r om t h e P o r tugue se N eo l i th ic s i te o f G ru ta d o Z ambu ja l , Me l ide s , h ave b een c ompa red t o Me so l i th ic s amp le s f r om t h e Muge m idd en s i te s o f Mo i ta d o S ebe s t iao a nd C abeco d aA r ruda . N o e f f ec t ive d i sc r im ina t ion o f t h e t empo ra l s amp le s w a s p o s s ib le , a nd c l u s te r ing t e chn ique s p roduced n o s i t e -cen te r ed g r oup ing s ( Me ik le john a nd S chen tag , n d ) . P a ra l le l f i nd ings h ave b een m ad e f o r t h e s ou th o fF rance b y D uday ( 1 977 ) , t h ough s amp le s i ze i s ap rob lem i nt h el a t te rc a se . C ong ruen tw i th s u ch c a se s a st h ea bove , i tc an b e d emon s t ra t ed t h a t t h e re i s c hange i n f o rm

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p o s i t ive p opu la t ion s f r om t h e e a s t . H oweve r , t h e o r ig in o ft h i s h ypo the s i s i sn o tb a sed o n e v idence o f t h e a n t iqu i ty o f t h e B a sque s , b u t o n t h e i r R h f r equenc ie s , t h u s c r ea t ing a c en t ra l c i r cu la r i ty w i th in t h e a r gumen t . L iv ing s tone f u r the r p o in t s o u t t h a t h i gh c de f r equen c ie s a r e a l so s e en i n o t he r a l p ine g roup s , s ugge s t ive o f ap o s i t ive s e lec t ion f o r t h i s a le le c omp lex u nde r t h i s e nv i ronmen ta l c i r cums tance . F u r the rmo r e , B an tu e xpan s ion i n s ub -Saha ran A f r i ca , am uch b e t t e r d o cumen ted a nd r e cen t e ven t , i sn o t m i r ro red b y t h eg ene t ic v a r ia t ion d a ta . 3 )Af i na l p o in t t h a t n eed s r a i s ing i st h a t t h e P r inc ipa l C omponen t s a na ly s i s s ugge s t s t h a t t h e i n i t ia l a g r icu l tu ra l c o lon i sa t ion h a s b een t h e m o s t i mpo r tan tp opu la t ion m ovemen t t oa f f e c tE u rop e , s i nce t h e F i r s t C omponen t , c on s t i tu t ing 2 7% o f t h e t o ta l v a r ia t ion , i sl i nked t o t h i s e ven t . T he i mp l ica t ion i st h a t e ven t s s u ch a st h e B ronze a nd l i on A ge e xpan s ion s , t h e R oman h egemony a nd t h eM ig ra t ion p e r iod a r ea l lo fl e s s s i gn if icance i n t h e f o rma t ion o f t h e c u r r en t E u rop ean g ene t ic m akeup . T h i s m ay b e s o , b u t i tl e ave s r oom f o r t h ough t . G iven t h e a bove , w e b e l ieve t h a t t h e h uman b i o log ica l e v idence f o r d em ic d i f f us ion a s t h e e xp lana t ion f o r a g r icu l tu ra l o r ig in s i s , a t b e s t , i n conc lu s ive ( s e ea l so S oka le ta l , 1 990 ) . T he c a se f o r t h e c o lon i sa t ion o f E u rope b y N ea rE a s te rn f a rme r sr e s t st o al a rge e x ten to n t h e i d en t i f ica t ion o f ad i e ta ry s h if t f r om f o rag ing t o f a rm ing w i th t h e a r r iva l o f t h e p u ta t ive n ew p opu la t ion . O s t eo log ica l d a ta p r e sen ted a bove , s u ch a st h er i se i nc a r ie s i nt h e Med i te r r anean l a te Me so l i th ic , s how s t h a t d i e ta ry c hange i sa l ready o ccu r r ing i n s ome p r e -Neo l i th ic c on tex t s . T h i s r ef l ec t s ac u l tu ra l s h if t i n s ub s i s tence p a t t e rn , d e t ec ted a r cha eo log i ca l ly , t owa rd s g r ea te r u s e o f v ege ta l f o od s ( a nd a l so a qua t ic r e sou r ce s ) i nt h e E u rop ean l a te Me so l i th ic . F i na l ly ,w i th t h ep o s s ib le e x cep t ion o fp a r t so f c en t ra la nd s ou th -ea s te rn E u rop e , t h ee v id ence f o r c on t inu i ty i n m a t e r ia l c u l tu r e a nd s e t t lemen t p a t te rn a r gue s a ga in s t t h e a pp l icab i l i ty o f t h e c o lon i sa t ion m ode l b eyond t h e E u ropean c on t inen ta l c o r e ( Zve leb i l , 1 986b ; Z ve leb i l a nd Z ve leb i l , 1 988 ) .

w i th in t h e E u ropean Me so l i th i c . R eg iona l p opu la t ions o f t h e Me so l i th i c a r e n o t u n i fo rm i n a ppea r ance , t h e d i f f e rence s b ecom ing m o r e m a rked i n l a t e Me so l i th i c t i me s . C hange s b e tween t h e Me so l i th ic a nd N eo l i th ic p opu la t ion s h ave t ob e a s se s sed a ga in s t s u ch ah e t e rogenou s b a ckg round . I n s umma ry , c hange s n o ted b e tween M e so l i th ic a nd N eo l i th ic c anno t b e u sed t o s u ppo r t a mode l o f p opu la t ion r e p la cemen t i fi t h a s e i the r b e en s hown t h a t s u ch c hange h a s a l r eady b een u nde r w ay i nt h e Me so l i th ic , o r i f t h e t y pe o f c hange i se qu iva len t t o d i f f e r ence s a l r eady p r esen t b e twe en Me so l i th i c p opu la t ion s . T he d emon s t ra t ion o f p opu la t ion c hange a t t h e M e so l i th ic -Neo l i th ic b ounda ry r e qu i r e s e v idence f o r amod if ica t ion i nt h e t r a jec to ry o fc hange , o r e v idence f o r d i s junc t ion i nt h e o ve ra l l p a t ter n o f m o rpho logy b e tween r e p la ced a nd r e p la c ing g r oup s . A c a se f o r s u ch m o rpho log i ca l d i s junc t ion b e tween Me so l i th ic a nd N eo l i th ic p opu la t ion s h a s b een m ade b y V en d l( 1 986 ) f o r c e n t ra l , a nd b y Menk a nd N eme ske r i ( 1 989 ) f o r s o u th -ea s t E u rope . H oweve r , t h e r e i s a n a l te r na t ive e xp lana t ion , w i th s u ch c hange b e ing s e en a st h er e su l to f ad i e ta ry s h i f tt o as o f te rd i e t , h i ghe r i n f i sh a nd /o r c a rbohyd ra te r e sou r ce s , r a the r t h an a s ar e su l t o fp opu la t ion r e p la cemen t ( y 'Edynak , 1 978 , 1 989 ; Z ve leb i l a nd Z ve leb i l , 1 9 88 ) . G ene t ic p a t t e rn ing o f e x tan t p opu la t ion s h a s b een u sed b y Amme rman a nd C ava l l i -Sfo r za ( 1 984 ) t o s uppo r t ac o lon i sa t ion m ode l f o r t h e d i f fu s ion o f f a rm ing i n E u rope , r e f e r r ed t o a s d em ic d i f fu s ion . A spe c t s o ft h e m ode l h ave b e en b o th s uppo r ted a nd c r i t ic i s ed b y a r chaeo log i s t s , p hy s ica l a n th ropo log i s t sa nd g ene t ic i s t sa l ike . We f i nd i ti n s t ruc t ive t h a t t h e re i s at e ndency f o r w o rke r s t oc r i t ic i se a spe c t s o f t h e s t udy r e levan t t o t h e i r o wn f i e ld o f e xp e r t i se w h i le a c cep t ing t h o se a spec t s i nf i e ld so t he rt h an t h e i ro wn . T hu s t h e a r chaeo log ica l a sp ec t s o f t h e v o lume h ave r e ce ived m o s t s uppo r t f r om p hy s ica l a n th ropo log i s t s a nd g ene t ic i s t s b u t n o t f r om a r chaeo log i s t s a nd v i ce v e r sa . We f i nd t h e f o l low ing p o in t s o fp a r t i cu la r i n te r e s t w i th r e ga rd t ot h eg ene t ic e v idence : 1 ) T he d i f fu s ion m ode l d o e s n o t d e r ive d i re c t ly f r om t h e r aw b l ood g r oup a nd p ro te in p o lymo rph i sm d a ta b u t f r om t h e P r in c ipa l C omponen t s d e r ived f r om t h e d a ta . A s Mo r ton ( 1 985 ) p o in t s o u t , b o th t h e m a thema t ica l b a s i s o f t h e c omponen t s a nd t h e i r i n te rp r e ta t ion i s h eu r i s t ic . T hu s t h ough s uppo r t f o r t h e d i f fu s ion m od e l c an b e f ound i nt h e P r inc ipa l C omponen t a na ly s i s , t h e h ypo the s i s d oe s n o t n ece s sa r i ly d e r ive f r om t h ea na ly s i s . 2 ) B o th Mou ran t ( 1 985 ) a nd L iv ing s ton e ( 1 985 ) p o in t o u t t h a t ak ey a s sump t ion o f t h e m ode l , i n c lud ing i n te rp re t a t ion o f t h e P r inc ipa l C omponen t s , i st h a t R h n ega t ive ( h omo zygo te s f o r t h e c de a le le c omp lex ) B a sque s r e p r e sen t a p r e -Neo l i th ic s u rv iva l , l a te r r e p la ced b y R h

D iscus s ion The I nd igenous a nd Co lon is t N eo l i th isa t ion i n Europe I no u rv i ew ,t h eb i o log ica ld a ta o u t l ined a bove a dd t oo t he re v idence t os how t h a t : 1 )T he a s sump t ion s m ade b y t h e c o lon i sa t ion o r d em ic d i f f u s ion m ode l m ay n o t b e f u l ly w a r ran ted , a nd m ay n o t n ece s sa r i ly f o l low f r om t h ee v idence p r e sen ted f o rt h em . 2 ) T he mod e l d oe s n o t a c coun t f o r t h e t r an s i t ion t of a rm ing i na r ea so fE u rop eo t he rt h an p o s s ib ly t h ee a s te rn Med i t e r ranean , s ou ther nI t a ly

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a va i lab le t oE u ropean l a te Me so l i th i c g r oup s u n t i l t h e 6 t h m i l lenn ium b e . I n s umma ry , t h e w e s t Med i t e r ranean p a t te rn w ou ld s ugge s t t ou s e a r ly s t age s i nt h e i n ten s ive u s e o fp l an t f o ods , a s y e t n o t a tended b y m a rke r s o f s t re s s a nd r e duced h ea l th w h ich a r e c ommon a mong m any p r eh i s to r ic a g r icu l tu r i s t s ( e .g . C ohen a nd A rme lago s , 1 984b ) .

a nd t h e D anube /Rh ine v a l ley s y s tem . O u t s ide o f t h i s c on t inen ta l c o r e , i tw ou ld a pp ea r l i ke ly t h a t f a rm ing w a s a dop ted b y r e s iden t Me so l i th ic p opu la t ion s ( Denne l l , 1 983 ; B a rke r , 1 985 ; Z ve leb i l , 1 986b ; P r ice , 1 987 ; Z ve leb i l a nd Z ve leb i l , 1 988 ) .The b i o log ica l e v idence i s c on sonan t w i th t h e n o t ion o f as l ow a nd g r adua l t r an s i t ion t of a rm ing i n mo s t a r ea s o f E u rope , a p ro ce s s w h ich w a s n o t c omp le t ed u n t i l t h e l a t e N eo l i th i c : as ugge s t ion m ade b y o ne o fu se a r l ie r o n t h e s t r eng th o f c u l tu ra l e v idence a l one ( Zve leb i l , 1 986a ,b ) . S uch c onc lu s ion s s h i f t o u r a t t en t ion t o t h e c ond i t ion o f t h e i n d igenou s c ommun i t ie s o f t h e Me so l i th i c . H e re w e h ave t o m od i fy o u r f i r s t p r opo s i t ion , c once rn ing t h e g ood h ea l th s t a tu s o f Me so l i th ic p opu la t ion s . C lea r ly , ad ua lp a t te rn o f p a tho logy i se me rg ing : a" we s t Med i te r ranean " z one , m a rked b y t h e h i gh i n c idence o fc a r ie s a nd t h er e la t ive a b sence o fo t he ri n su l t s ,a nd t h eo t he r , " no r th -ea s t E u rop ean " o ne , m a rked b y t h e o ccu r en ce o fi n d ica to r so fn u t r ien td e f i c iency a nd b y t h ea b sence o fc a r ie s( F ig . 1 ) .

L a te Meso l i th ic : T he Nor th a nd E as t European P a t tern I ft h ep a tho logy p a t t e rn o b se rved a tI r on G a t e s a nd i nt h e s o u th B a l t ic i sr e p re sen ta t iv e o f o t he r h un te r -f i she r g r oup s i n n o r the rn a nd e a s te rn E u rope , t h en t h e se l a te Me so l i th ic c ommun i t ie s w e r e w o r se o f fi nt e rm s o fh ea l th t h an w e r e t h e i r w e s t Med i te r rane ran c oun te rpa r t s . T h i s a ppa ren t p a radox m ay p o se ac ha l lenge f o rt h o se w ho v i ew f o rage r g r oup s d ependan t o n a qua t i c r e sou r ce s a s " a f f luen t " ( i .e . Wana tab e , 1 968 ; S u t t le s , 1 968 ; A kazawa , 1 986 ; H ayden e ta l , 1 985 ) . H oweve r , t h e r e m ay b e g ood r e a sons f o r t h e h i ghe r i n c iden ce o f s t r e s s a mong t h e h un te r -f i she r g r oup s i n n o r the rn a nd e a s te rn E u rop e . E ven t h ough , a ts i t e s s u ch a s V la sa c , t h e r e i se v idence f o r t h e u s e o fp l an t f ood s ( P r in tz , 1 987 ) , a b sence o f c a r ie s , a nd a h i gh i n c id ence o f c a lcu lu s a nd e name l h ypop la s ia s ugge s t ap ro te in d om ina ted d i e t a nd r e cu r r ing n u t r ien t s t re s s . L a ck o f a r e gu la r c a rbohyd ra te d i e t m ay h ave b een a c on t r ibu to ry f a c to r i n t h e f a i lu re t o p r even t p e r iod ic s ho r t age s a r is ing f r om s e a sona l f l uc tua t ion i n r e sou r ce s i n t h e E u ropea n c on t inen ta li n t e r io r . B u t t h e r e i sa n a l te rna t ive , a nd i no u r v i ew m o r e c onv in c ing e xp lana t ion : t h a t i n c r ea sed r e s iden t ia l p e rmanence a nd d ependance o n af i sh d i e t r e su l ted i n p a ra s i t ic i n f e s ta t ion s a nd i n f ec t ion s w h ich p r ed jud iced t h e h ea l th o f t h e se h un te r f i she r c ommun i t ie s a nd r e su l t ed i n t h e o b se rved r a nge o f p a tho log ica l c ond i t ion s . E v id ence f r om an umbe ro fw ide rang ing c on tex t s o u t s ide E u rop e s uppo r t s t h is h ypo the s i s : a b sence o fc a r ie s , h i gh i n c iden ce o fr i cke t s , o s teoma la c ia , a nd p o ro t ic h ype ro s to s i s m a rked t h e p a t te rn o f p a tho logy s e en , f o r i n s tance a mong t h e f i sh e a t ing c ommun i ty a tR a t sa l -Ham ra a l ong t h ee a s t A rab ian c o a s t ( Ma cch ia r e l l i , 1 989 ) . K en t ( 1 986 ) n o te s h i gh i n c iden ce o f p o ro t ic h ype ro s to s is a mong t h e E sk imo a nd a mong t h e p r eh i s to r ic m a r ine -dependan t C a l i fo rn ian I nd ian s . I n c rea sed s e den t i sm a nd d ep endance o n a qua t ic r e sou r ce s i s ac ommon d enom ina to rf o ra l t h e se g r oup s . H ow m uch s t re s s i s r e p resen ted b y t h e se c ond i t ion s ? H ow f a r c an s t r e s s b e s e en a s a n i n cen t ive f o r t h e a dop t ion o f f a rm ing? T he re a r e w ide ly d i ve rgen t v i ew s o n t h i s m a t te r . I n r e f e r ence t o t h e L evan t , S m i th e t a l ( 1 984 ) c ommen ted :

L a te Meso l i th ic : The Wes t Med i terranean P a t tern T he w e s t Med i te r ranean p a t t e rn , m a rked b y c a r ie s , i s c oeva l w i th a n i n c r ea sed i n take o f s t a r chy c a rbohyd ra te f o od s , r e su r r ec t ing t h e q ue s t ion ,p o sed 1 5 y ea r s a go b y C la rke ( 1 976 ) o f p l an t h u sbandry i n t h e Me so l i th i c o f t h i s a r ea ( t h i sp a t ter nm ay a ppea re a r ly i nt h e Me so l i th ic a s s e en f r om t h e e v idence o f s i te s s u ch a s U zzo ( Bo rgogn in i -Ta r l i e ta l , 1 989 ; C on s tan t in i , 1 989 ) a nd E lC ing le V e rme i l ( Tu rbon , 1 986 , 1 989 ) . Y e t , c ompa r a t ive d a ta f r om k nown f a rm ing c on tex t s d o n o t r e p l ica te t h i s w e s t Med i te r ranean p a t te r n . T he L evan t ine s i tua t ion s how s al e s se r i n c idence o fc a r ie s i nt h e N a tu f ian a nd t h e N eo l i th ic , b u t a g r ea te r i n c id ence o f o t he r s t r e s s r e la ted i n su l t s , s u ch a s c r ib r a o rb i ta l ia , p o ro t ic h ype ro s to s i s , e name l h ypop la s ia a nd o s teopo ro s i s ( Ange l , 1 984 ; S m i th e ta l , 1 984 , 1 989 ) , s how ing t h a t , o n t h e w ho le , t h e L evan t ine p opu la t ion s w e r e l e s s h ea l thy t h an t h e w e s t Med i te r ranea n o ne s . N a t ive N o r th Ame r i can c u l t iva to r s s u f f e r ed f r om h i gh l e ve ls o f b o th c a r ie s a nd o t he r i n d ica to r s o f n u t r i t iona l s t re s s ( Ca s s idy , 1 984a ,b ; C ook , 1 984 ; G oodman e ta l , 1 984a ; L a r sen , 1 984 ; P e rz ig ian e t a l , 1 984 ; R o se e ta l , 1 984 ) w i th t h e d i s t inc t ion t h a t , u n l ike t h e w e s t Med i te r ranean p a t te rn ,c a r ie s a l so o ccu red a mong t h e c h i ld r en . T h i s s ugge s t s t h a t s o f tc a rbohyd ra te d i e t w a s f e d t oj u ven i le s i n N o r th Ame r i ca , ap a t te rn t h a t s e em s t ob e a b sen t i nE u rope ( Me ik le john e ta l , 1 988 ) ( a tt h e s i t e so f Mo i ta d o S eba s t iao a nd C abe co d a A r ruda t h e r e a r e n o c a se s w he re c a r ie s c an b e d emon s t ra t ed p r io r t ot h e e r up t ion o f t h e t h i rd m o la r s a tc a 1 8 y ea r s o fa ge ) . T h i s m ay r ef le c t t h e u s e o fp o t t e ry a mong t h e N o r th Ame r ican l a te Wood land a nd M i s s i s s ipp ian g r oup s f o r p r epa r ing s o f t m ea l s ( Bu ik s t ra e t a l , 1 986 ) , a n o p t ion t h a t w a s n o t

" The i n i t ia l s t ep s i n p l an t a nd a n ima l d ome s t ica t ion w e r e n o t a s soc ia ted w i th

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e nv i ronmen ta l h ea l th s t a tu s " .

s t r e s s

o r

d e t e r io ra t ing

A s im i la r p o s i t ion h a s b een a dop ted b y a n umbe ro fo t he rr e sea r che r s ( i . e . R oo seve l t , 1 984 ; B u ik s t r a e t a l , 1 986 ) , w h i le o t he r w o rke r s m a in ta in t h a t p opu la t ion p r e s su re a nd r e sou r ce s t r e s s w e re t h e p r inc ipa l c au se s u nde r ly ing t h e a g r i cu l tu ra l t r an s i t ion ( i . e . C ohen , 1 977 ; B in fo rd , 1 9 83 ; R edd ing , 1 988 ) . T he D e mography o ft he Meso l i th ic N eo l i th ic T rans i t ion D emog raph ic p a rame te r s s hou ld p l ay am a jo r r o le i n t h i s d i scu s s ion , s i nce s u r v iva l a nd r e p roduc t ion a r e u l t ima t e m ea su r e s o ff i t ne s s a nd ac ond i t ion f o r s o c ia l , a s w e l l a s b i o log i ca l , r e p roduc t ion . B o cque t -Appe l a nd Ma s se t ( 1 982 ) a nd J ohans son a nd H o row i tz ( 1 986 ) h ave o u t l ined p r ob lems i n he ren t i n t h e p a la eodemog r aph ic a pp roach . S uch p rob lem s ( r ep re sen ta t ivene s s o f t h e s amp le , a s sump t ion s o f c l o su r e a nd c on s tan t g r ow th r a te , e t c . : s e e G r aue r , t h i s v o lume ) h ave l e d , f o r e xamp le , t o t h r ee d i f f e r en t e s t ima te s o f l i f e e xpe c tancy a t V la sa c ( Neme ske r i , 1 978 ; y ' Edynak , 1 989 ) . G iven t h i s s i tua t ion , i tf o l low s t h a t a ny p a laeodemog raph ic a s se s smen t h a s t ob e t r ea ted w i th c au t ion . H oweve r , a t av e ry g ene ra l l e ve l , t h e r e s e ems t oh ave b een a s l ow i n c rea se i nl i f e e xpec ta ncy t h roughou t t h e P a laeo l i th ic t or e ach t h e v a lue o f 3 0 -34 y ea r s i n t h e l a te U ppe r P a laeo l i th ic a nd M e so l i th ic o f E u rope a nd t h e N ea r E a s t ( Con s tand se - We s te rmann e t a l , 1 984 ; A nge l , 1 9 84 ) . I n t h e N eo l i th ic , l i f e e xpe c tancy v a lue s r ema ined s t ab le , o r h ave d ec l ined s l igh t ly i n E u rope a nd t h e N ea r E a s t ( Neme ske r i , 1 978 ; A nge l , 1 984 ) , w h i le i n N o r th Ame r ica l i f e e xpe c tancy o f a g r i cu l tu ra l p opu la t ion s i s o f ten r e du ced d r ama t ica l ly c ompa r ed t o h un te r g a the r e r s a nd t h e t r an s i t iona l h un t ing -cu l t iva t ing c ommun i t ie s ( J ohan s son a nd H o row i tz , 1 986 ; B u ik s t ra e ta l , 1 986 ; E i senbe rg , t h i s v o lume ) . A t t h e s ame t i me , r e p roduc t ive r a te s o f p r eh i s to r ic f a rme r s r ema ined g ene ra l ly u na f f ec ted a nd t h e r e i se v idence o fp opu la t ion i n c r ea se . T h i s p o in t s t o t h eo b se rva t ion m ade b yR indo s( 1 985 ) : i n c r ea sed f i t ne s s i sn o t a lway s t h e s ame a s a dap t ivene s s . W i th t h e t r an s i t ion t of a rm ing , i n c r ea sed f i t ne s s , i nt e rms o fr e p rodu c t ion , h a s b een a ta ined a tt h e e xpense o f a dap ta t ion : t h a t i s , o f e qu i l ib r ium b e tween t h e p opu la t ion a nd t h e e nv i ronmen t , r e su l t ing i nd ec l ine i nl i f ee xpe c tancy ,d e c r ea se i n s t a tu re a nd i n c r ea se i ns t re s sd u r ing t h eN eo l i th ic . I ti so n ly i ne x t r eme c a se s , w hen s t r e s s b ecome s a cu te a nd t h rea ten s t h e s u rv iva l o f ac ommun i ty t h a t f i tne s s w i l l d ec l ine w i th m a ladap ta t ion , a s w a s t h e c a se w i th t h e N o r se s e t t lemen t i n G r een land , o r w i th t h e A ve rbu ch M i s s i s s ipp ian c ommun i ty i nN o r th Ame r ica ( s ee E i senb e rg , t h i s v o lume ) . A ga ins t t h i s b a ckg round , i ts e ems t h a t l e ve l s o f s t res s i n t h e Me so l i th i c d i d n o t r e a ch t h e

t h r e sho ld w h ich w ou ld t h rea ten t h es u rv iva lo ft h e c ommun i t ie s i nq ue s t ion o rc au se ad e c l ine i nl i f e e xp ec tancy . E ven a t V la sa c , N eme sk e r i a nd L engye l ( 1 978 ) a c co rd t h e p opu la t ion " a r a the r g ood h ea l th s t a tu s " , w h i le i t s u nco r rec ted l i f e e xpe c tancy i se s t ima ted a t3 6 y ea r s ( Neme ske r i [ 1 978 ] : o t he re s t ima te sr a nge f r om 2 6t o4 4 y ea r s , s e e a l so y ' Edynak [ 1 989 ] ) . L i t t le c hange i nt h e se p a rame te r sc an b es e en a s ar e su l to ft h e t r an s i t ion t o t h e N eo l i th ic : o n t h e p r e sen t e v idence , a s l igh t ly h i ghe r e v idence f o r p a tho logy i n t h e Me so l i th i c ( s ee a bove ) i sc ompen sa t ed f o rb y t h e s l igh t ly l owe r l i f e e xpec tancy e s t ima te s i n t h e N eo l i th i c . T h i s w ou ld s uppo r t t h e n o t ion o f a g r adua lt r an s i t ion t of a rm ing i nE u rope . Wh i le t h e se c on s ide ra t ion s s uppo r t t h e v i ew t h a t l e ve l s o f p a tho logy a nd s t re s s i n t h e Me so l i th i c w e re n o td e t r imen ta l t ot h e s u rv iva lo f ac ommun i ty , t h ey d o n o t e xc lude t h e p o s s ib i l i ty t h a t t h e l e ve l so fs t re s s , s u ch a st h ey w e re , s e rved t oe ncou rage t h e a dop t ion o f f a rm ing . I ns e ek ing t o u nde r s tand t h i s p ro ce s s , w e a ga in h ave t o d i s t ingu i sh b e twe en t h e d i f f e ren t p a t te rns o f t h e t r an s i t ion i n t h e s ou th -we s t a nd i n t h e n o r the rn a nd e a s te rn p a r t so fE u rope . T he Agr icu l tura l T rans i t ion i n t he Wes t Med i terranean I n t h e w e s t Med i te r ranean , t h e f i r s t e l emen t s o f a g ro -pa s to ra l " Neo l i th ic " f a rm ing w e re a dop t ed r e la t ive ly e a r ly , a ta bou t6 000 b c , b u t t h e a dop t ion w a s s e le c t ive a nd s t agge r ed o ve r t wo m i l lenn ia , i n t h a t d i f f e r en t e l emen t s o f t h e s y s t em : c ap r ine s , b ov ine s , c e r ea l s , p u l se s , c e r am ic s a nd f a rm ing t e chno logy w e r e a dop t ed s e pa r a te ly a nd g r adua l ly a c ro s s t h e r e g ion ( Gu i la ine , 1 987 ; V ague r , 1 989 ; B inde r , 1 989 ; L ew thwa i te , 1 986 ; Z ve leb i l a nd Z ve leb i l , 1 988 ) . I n o t he r w o rd s , w h i le t h e s p r ead o f s ome N eo l i th i c t r a i t s w a s r a p id t h roughou t t h e Med i te r r anean ( L ew thwa i t e , 1 986 ) , t h e s ub s t i tu t ion o f t h e l o ca l b y t h e N eo l i th ic s y s t em w a s l o ng . I n o u r v i ew , t h i s p a t te rn i sc on sonan t w i th t h e n o t ion t h a t h u sband ry p r ac t ice s w e re n o t r e a l ly n ew : t h ea dop t ion o fn eo l i th ic c u l t igen sa nd d ome s t ica te s r e p re sen t ed a c on t inua t ion o f p r ac t ice s t h a t w e r ea l r eady p r e sen t , i nt h e f o rm o f a r bo r icu l tu re o ff i g s , o l ive s a nd n u t -bea r ing t r ee s , a nd p e rhap s i nt h e f o rm o f c u l t iva t ion o f s ma l l s e eded g r a s se s a nd p u l se s . G iven t h i s s i tua t ion , a nd b ea r ing i nm ind t h e l ow l e ve l so fs t r es s n o t ed i nt h e We s t Med i te r ranean , w ed on o tb e l ieve t h a t a dop t ion o f f a rm ing w ou ld c onf e r a n a dvan tage o n h ea l th g r ound s a l one . Wh i le l e ve l s o f s t r e s s d e t ec ted i n Me so l i th ic p opu la t ion sm ay h ave b e en ac on t r ibu t ing f a c to r i nt h e a dop t ion o f f a rm ing , o t he r m o r e i mmed ia te c au se s m u s t b e s ough t w i th in t h e s o c io - e conom i c o rgan i sa t ion a t a r e g iona ll e ve l .

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C onc lu s ion

T he Agr icu l tura l T rans i t ion i n Nor thern a nd E as tern Europe

I n s umma ry , o u r s u rvey o f p a tho logy i n E u ropean o s t eo log ica l s amp le s r e la ted t o t h e t r an s i t ion t o f a rm ing r e vea led u nexp ec t ed v a r iab i l i ty i n t h e i r h ea l th s t a tu s , r e la ted t o d i f f e r en tp a t t er n si nt h e i rd i e ta nd l i f e s ty le . O nt h e w ho le , w e f e e l t h a t t h e l e ve l s o f s t r e s s o b se rved w e r e n o ts u f f i c ien tt or e ga rd t h i s s t re s sa s ad i re c t c au se o f t h e a gr icu l tu ra l t r an s i t ion . I mmed ia te c au se s o f t h e t r an s i t ion mus t b e c on t ingen t o n r e g iona l s o c io -e conom ic a nd e co log ica l c ond i t ion s . I n t h e w e s t Med i t e r ranean , t h e a dop t ion o fN eo l i th i c c u l t igens a nd d ome s t ica te s c an b e s e en a s ad eve lopmen t o f p a t te rn s o f h u sband ry a l r eady p r e sen t i n t h e Me so l i th ic ( i e . C la rk e 1 9 76 ) . I n n o r ther n a nd e a s te rn p a r t s o f E u rop e , s o c ia l c ompe t i t ion , f o s te r ed b y t h e p r ox im i ty o f f a rm ing c ommun i t ie s , m ay h ave p r ov ided a n i mmed ia te c au se f o r t h e a g r icu l tu ra l t r an s i t ion . S o c ia l a nd e co log ica l e xp lana t ion s f o r t h e t r an s i t ion t of a rm ing h ave b een o f ten t r ea ted a s m u tua l ly i n compa t ib le , b u t t h i s d oe s n o t n eed t o b et h ec a se . B o th s o c ia la nd e co log ica lc ond i t ion s m ay h ave a c ted i nc omb ina t ion t ob r ing a bou t t h e a dop t ion o ff a rm ing ; s i nce i nb o th c a se s t h er a nge o f s o lu t ion s t o p rob lems c au sed b y t h e se c ond i t ion s w e r e l o ca t ed w i th in t h e h un te r g a the r e r s o c ie t ie s . F u tu re r e sea r ch s hou ld e xam ine s o c ia l mode l s f o r t h e t r an s i t ion t o f a rm ing w i th in t h i sf r amewo rk . T he b i o cu l tu ra l s t udy o f h uman s k e le ta l r ema in s , a s a r gued f o r h e r e , i ss t i l l i ni t s e a r ly s t age s f o r E u ropean S t one A ge p opu la t ion s . T he i n f e r ence s f r om t h e b ody o f d a ta c u r r en t ly a va i lab le a r ec omp rom i sed b y s ma l l s amp le s i ze s , l a ck o fr e p r e sen ta t ivene s s , a nd al im i t ed r a nge o f a na ly s i s . R a r e ly d o t h e c u l tu ra l a nd p a tho log i ca l /d emog raph ica l a na ly se s o f t h e s k e le ta l p opu la t ion s c o in c id e . T hus f o r i n s tance , t h e n umbe r o f b u r ia l c on tex t s a na ly s ed f o r p a tho logy i n t h e w e s t Med i te r ranean i s b u t a s ma l l s amp le o f t h o se i n e x i s tence ( Tu rbon , 1 989 :f ig .1 ; G ana lda , 1 989 ) . O r , t h e a na ly s i s o f m o r tua ry v a r ia t ion a tV la sa c , a nd t h e r e la t ion sh ip b e tween s o c ia l s t a tu s a nd p a tho logy a t V la sac w ou ld b eo fg r ea ti mpo r tance f o rs oc ia lm ode l so f t h e a g r i cu l tu r a l t r an s i t ion r e f e r red t o a bove . Mo r eove r , V la sa c i s n o t t h e o n ly s k e le ta l p opu la t ion i n t h e h on G a te s , a l though i ti st h e o n ly o ne c omp r ehen s iv e ly a na ly sed ( Ga r a shan in , 1 978 ) . F u l l a na ly s i s o ft h e s k e le ta l r ema in s f r om L ep en sk i V i r , C u ina T u r cu lu i , a nd P ad ina w ou ld a dd c r u c ia l i n f o rma t ion t o o u r u nde r s tand ing o f t h er e g ion . S im i la r ly , t h ea na ly s i s o ft h e m o r tua ry v a rab i l i ty a t O leneo s t rov sk i Mog i ln ik ( O 'Shea a nd Z v e leb i l , 1 9 84 ) , t h e l a rge s t Me so l i th i c s i t e i n E u rope , h a s n o t b e en m a t ched , t o d a t e , b y p a la eopa tho log ica l s t ud ie s , a t l e a s t n o t i n p ub l i shed f o rm . T he p auc i ty o f f uly a na ly sed N eo l i th i c s amp le s h a s a l ready b een m en t ioned . F i na l ly ,t h ec ompa r ab i l i ty o fd i f f e ren tc a se s t ud ie s

I n t h e s ou th B a l t i c a nd t h e I r on G a te s , t h e s i tua t ion w a s d i f f e r en t . T he r e , a g ro -pa s to ra l f a rm ing w a s a dop ted r e la t ive ly l a t e b y h un t e r f i she r g roup s , a f te r i t h ad b e en a va i lab le i n a d ja cen t r e g ion s f o r a bou t 1 000 y ea r s ( Zve leb i l a nd R ow ley -Conwy , 1 984 , 1 986 ; K oz low sk i a nd K oz lowsk i , 1 986 ) . D u r ing t h i s " a va i lab i l i ty p ha se " , e xchange c on ta c t s d eve loped b e tween f o rage r s a nd f a rme r s a c ro s s t h e a g r icu l tu ra l f r on t ie r( B lankho lm , 1 987 ;V oy t ek a nd T r ingham , 1 989 ; C hapman , 1 989 ) . When t h e a c tua l t r an s i t ion o cu r red ( c a . 3 100 b e i n D enma rk , c a . 4 500 b c i n t h e I r on G o rge s ) , i to ccu r ed r a p id ly , w i th in a p e r iod o f p e rhap s 2 00 y ea r s . I n o t he r w o rd s , a nd i nc on t ra s t t ot h e w e s t Med i te r ranean , t h e a va i lab i l i ty p ha se w a s l o ng , b u t t h e s ub s t i tu t ion p ha se w a s s ho r t ( Zve leb i l a nd R ow ley -Conwy , 1 984 , 1 986 ) . T h i s w a s t h e c a se d e sp i te m o r e e v iden ce f o r s t re s s p r e sen t i nt h e se c ommunn i t ie s . Wa s t h i s d ue t o t e cho log ica l a nd s o c ia l i n ve s tmen ti nt h es p e c ia l i sed ,a qua t ica l ly -o r ien ted e conomy ( Akazawa 1 986 )? Wa s t h i s b ecau se e xchange o f p roduc t s w i th t h e f a rme r s , o r p e rhap s , i n t h e c a se o f t h e h on G a t e s , l i m i t ed p l an t h u sband ry , m ay h ave g one s ome w ay t o b uf f e r t h e se c ommun i t ie s a ga in s t n u t r i t iona l s t re s s ? Y 'Edynak ( 1 989 ) n o te s s ome d ec l ine i n e name l h ypop la s ia i n t h e l a te r l e ve l s a t V la sa c , a nd l i nk s t h i s w i th g r ea te r a va i lab i l i ty o f p l an t f o od ( s ee a l so P r in tz , 1 987 ) . B o th o ft h e se f a c to r s m ay h ave p l ayed a p a r t -b u t t h e d e lay i n t h e a dop t ion o ff a rm ing ,w h i le s t r e s sw a si ne v idence , d oe sn o ts uppo r tt h ev i ew t h a tr e s idua ls t re s ss u ch a s w a s o b se rved i ne a s t er n a nd n o r the rn E u rop e c an p rov ide t h e m a in r e a son f o r t h e a g r icu l tu ra l t r an s i t ion . I n ev i tab ly ,s u ch ac onc lu s ion i sb ound t os h i f t a t t en t ion t os o c ia l f a c to r s . S oc ia l c ompe t i t ion a s t h e a l ter na t ive e xp lana t ion f o r t h e t r an s i t ion t o f a rm ing h a s b een a dvanced b y Mad sen ( 1 982 ) a nd J ennbe r t ( 1 984 ) f o r s ou the rn S cand inav ia , b y B ende r ( 1 985 ) f o r B r i t tany a nd b y L ew thwa i te ( 1 986 )f o rt h ew e s t Med i te r ranean . I n t h i s p ape r , w e h ave a r gued f o r i n c r ea se i n s o c ia l d i f f e r en t ia t ion , s eden t i sm a nd p opu la t ion d en s i ty i n t h e l a t e Me so l i th ic i n E u rop e , i n p a r t icu la r a mong t h e h un t e r -f i she r c ommun i t ie s o f n o r the rn a nd e a s ter n E u rope . S o c ia l c ompe t i t ion , i n w ha teve r f o rm , m ay h ave b e en g i ven a n a dded i mp e tu s b y al o ng t r ad i t ion o f e xchange i ne xo t ic g ood s a c ro s s t h e a g r icu l tu r a l f r on t ie r , t h e e v idence f o r w h ich c an b e s e en i n c e r am ic s a nd s t one a r te f a c t i mpo r t s a mong t h e h un te r f i she rc ommun i t ie so fS cand inav ia a nd t h e I r on G a te sr e g ion . I ts hou ld , f i na l ly , b e n o t ed t h a t l o ng d i s tance e xchange i ne xo t ic g ood s c an a l so b e d emon s t ra t ed a moung t h e Me so l i th i c c ommun i t ie s t h em se lve s ( Con s tand se We s t e rmann a nd N ewe l l , 1 988 ) .

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B inde r ,D .( 1989 ) . A spec t sd el a Wo l i th isa t ion d an sl e s A i re s P adane , P rovenca le e t L igu r e . I n N eo l i th i sa t ion s e d s . 0 . A u r enche a nd J .C auv in . p p . 1 99 -225 . B r i t i sh A r chaeo log ica l R epo r t s I n t er na t iona lS e r ie s5 16 . B inf o rd , L .R . ( 1 968 ) . P o s t -P le i s tocene a dap ta t ion s . I n N ew P e r spec t ives i n A rchaeo logy . e d s . S . B in fo rd a nd L .R .B in fo rd .p p . 3 13 -341 .C h icago : A ld ine . B inf o rd L .R . ( 1983 ) . I n P u r su i t o ft h e P a s t . L ondon : T hame sa nd H ud son . B lan l cho lm , H .P . ( 1 987 ) . L a te Me so l i th ic h un t e r g a the re r s a nd t h e t r an s i t ion t o f a rm ing i n s ou the rn S cand inav ia . I n Me so l i th ic N o r thwes t E urope : R ecen t T rend s . e d s . P . R ow ley -Conwy , M . Z ve leb i l & H .P . B lan I cho lm . p p . 1 55 -162 . S heff ie ld : D epa r tmen to fA rcha eo logy a nd P reh i s to ry . B o cque t -Appe l , J . P . a nd Ma s se t , C . ( 1 982 ) . F a rewe l lt o p a laeodemog raphy .J .H um .E vo l . 1 :321 -333 . B og i t sh , B .J . a nd C heng , T .C . ( 1990 ) . H uman P a ra s i to logy . P h i lade lph ia : S aunde r s C o l lege P ub l i sh ing . B o lomey , A . ( 1 973 ) . A n O u t l ine o fl a t e E p ipa la eo l i th ic E conomy a t t h e " I ron G a te s " : T he E v id ence o n B one s .D ac ia 1 7 :112 . B o rgogn in i -Ta r l i , S .M . a nd R epe t to , E . ( 1985 ) . D ie t , d en ta l f e a tu r e s a nd o r a l p a tho logy i nt h e Me so l i th ic s amp le s f r om U zzo a nd Mo la ra c ave s ( S ic i ly ) . I n P ape r s i n I t a l ian A rchaeo logy I V ( I I ) . P reh i s to ry . e d s . C . Ma lone a nd S .S t odda r t . p p . 8 7 -100 . B r i t i sh A rcha eo log ica lR epo r t sI n te rna t iona l S e r ie s2 44 . B o rgogn in i -Ta r l i , S .M . D e l la S an t ina , D . F ranca lacc i , P .&R ep e t to ,E .( 1 989 ) .R econ s t ruc t ion o f Me so l i th ic d i e t u s ing d en ta l m ic rowea r a nd t r a ce e l emen t a na ly s i s . T he c a se o f G ro t ta d e l l 'Uzzo ( S ic i ly ) . I n P eop l ea nd C u l ture i nC hange .e d . I .H e r shkov i tz .p p . 2 83 -320 . B r i t i sh A rchaeo log ica l R epo r t s I n t er na t iona lS e r ie s , 5 08 . B r inch P e t e r sen , E . ( 1 973 ) . A s u r vey o f l a te P a laeol i th ic D enma rk . I n T he Me so l i th ic i nE urope . e d . S .K . K oz low sk i . p p . 7 7 -127 . Wa r saw : Wa r saw U n ive r s i ty P re s s . B r inch P e te r sen , E . ( 1 981 ) . B le v d e m yrde t a l le t r e? C openhagen : Munk sgaa rd . 3 1 p p . . [ S tena lde r e t j aeger f o lk f o r7 000 ä rs i den , 3 ] B u ik s t ra , J . E . K on ig sbe rg , L .W . a nd B u l l ing ton , J . ( 1 986 )F e r t i l i ty a nd T he D eve lopmen t o fA gr icu l tu re i nt h eP reh i s to r ic M idwe s t . Am .A n t iq . 5 1(3 ) :528 -546 C a lcagno ,J .M . a nd G ib son , K .R . ( 1 988 ) . H uman d en ta l r e duc t ion : n a tu ra l s e lec t ion o r p robab le mu ta t ion e f f ec t . Am .J .P hys .A n th ropo l . 7 7 (4 ) :505 -517 . C a rcu ima ru , M . ( 1 978 ) . L 'Ana ly se p ol in ique d e s c op ro l i the sd e l as t a t ion a r chaeo log ique d e V la sac . I n V la sac : A Meso l i th ic S e t t lemen t i n t h e I r on G a te s , v o l . I . e d . M . G a ra shan in . p p . 3 1 -34 . B e lg rade : S e rb ian A cademy o fS c ience s a nd A r t s . C a s s idy , C .M . ( 1 984a ) . N u t r i t ion a nd h ea l th i n a g r icu l tu ra l i s t s a nd h un te r -ga the re r s . I n N u t r i t iona l A n th ropo logy . e d s . N .W . J e rome , G .H . P e l to & R .F . K ande l . p p . 17 145 . P l ea san tv i l le , N ew Y o rk : R edg rave P ub l i sh ing C ompany . C a s s idy , C .M . ( 1 984b ) . S ke le ta l e v idence f o r p r eh i s to r ic s ub s i s tence a dap ta t ion i nt h eC en t ra lO h io R ive r V a l ley . I n P a leopa tho logy a t t h e O r ig ins o f A gr i cu l ture . e d s . M .N . C ohen a nd G .J . A rme lago s . p p . 3 07 -345 .O r lando : A cadem ic P re s s . C hagnon , N .A . ( 1 972 ) . T r iba l s o c ia l o rgan i sa t ion a nd g ene t ic m ic rod i f f e ren t ia t ion . I n T he S t ruc ture o f H uman P opu la t ion s . e d s . G .A . H a r r i son a nd A .J .

i sa dv e r se ly a f f ec ted b y v a r ia t ion i n a pp roa ch , c r i te r ia u sed i n i d en t if i ca t ion , a nd a n a l t o o f r equ en t l a ck o f a t ten t ion t o c on tex tua l i n fo rma t ion , b o th c u l tu r a l a nd e nv i ronmen ta l ( t he M e so l i th ic o f w e s t e rn a nd c en t ra l E u rop e i st h e o n ly p e r iod t o h ave h ad a c omp r ehen s ive e va lua t ion o f t h e c on t ex t o f a t t r ibu ted s k e le ta l r ema in s [ Newe l l e ta l , 1 979 ] ) . We h ave c ho sen t o i g no r e t h ese p rob lems i nt h i s s t udy , i nt h e h ope t h a t s i gn if ican t p a t ter n s m ay b ecome a ppa r en t n eve r the les s . D ue t o t h e se p rob lems , h oweve r , o u rc onc lus ion s mu s to fn ece s s i ty b ep r e l im ina ry , a nd w i l l p robab ly c hange a sn ew e v iden ce c ome s t ol i gh t . We f e e l t h an m any o f t h e p rob lem s o u t l ined a bov e c an b e o ve r come w i th in t h e i n t eg ra ted , c on t ex tua l , b i o cu l tu ra la pp ro a ch . I nt h i ss t udy , w e h ave t r i ed t oc omb ine t h ee x i s t ing b i o log i ca l d a ta a r i s ing f r om t h e s t udy o fh uman s ke le ta l r ema in s w i th t h e c u l tu ra l e v idence . P roce s se s o fr e cove ry a nd e x t rapo la t ion f r om m ode rn c a se s t ud ie s i n t roduce b i a sa nd u nce r ta in ty i nb o th s e t so fd a ta ( Con s tand se - We s te rmann e ta l , 1 984 ; Me ik le john e ta l , 1 984 ; J ohan s son a nd H o row i tz , 1 986 ) . T he r e la t ionsh ip b e tween t h e t wo i so f ten s ub t le a nd c omp l ica ted . Y e t t h e m ean ing o f e a ch i sm uch e nhanced b y t h e r e f e r ence t o t h e o t he r . I ft h i s p ap e r h a s s u cceeded i n d emon s t r a t ing t h e p o t en t ia l a nd t h e v a lue o f t h e b i o cu l tu ra l a pp roach , t h en i tw i l lh ave s e rved i t sp u rpo se .

R e f erence s A l caz awa , T . ( 1 986 ) . H un te r -ga the re r a dap t ion s a nd t h e t r an s i t ion t of o od p roduc t ion i nJ apan , I n H un te r s i n T rans i t ion . e d . M . Z ve leb i l . p p . 1 51166 . C amb r idge : C amb r idge U n ive r s i ty P r e s s . A lexande r sen , V . ( 1 988 ) . D e sc r ip t ion o f t h e h uman d en t i t ions f r om t h e L a te M e so l i th ic g r ave -f ie ld s a t S ka teho lm , S weden . I n T he S ka teho lm P ro jec t . I . Man a nd E nv i ronmen t . e d . L .L a r sson . p p . 1 06 -163 . S t ockho lm : A lmqv i s ta nd W ik se l l . A lexande r sen , V . ( 1988 ) .T he L a te -Me so l i th ic d en t i t ion i ns ou the rn S cand inav ia . R i v i s ta d iA n tropo log ia , 6 6 ( S upp lemen t ) ,p p . 1 9 1-204 . Amme rman , A . J . a nd C ava l l i -Sf o rza , L .L . ( 1 984 ) . T he N eo l i th ic T ran s i t ion a nd t h eG ene t ic s o fP opu la t ion s i nE urope .P r inc e ton : P r ince ton U n ive r s i ty P re s s . A nd e r sen , S .H . ( 1987 ) . T yb r ind V ig : A s ubme rged E r teb0 1 1e s e t t lemen t i n D enma rk . I n E u ropean W e t land s i n P reh i s to ry . e d s . J .M . C o le s a nd A .J . L awson . p p . 2 53 -280 .O xf o rd : C la rendon P r e s s . A nde r son , R .M . a nd May , R .M . ( 1 979 ) . P opu la t ion b i o logy o fi n f ec t iou sd i sea se s .N a ture ,2 80 :361-367 . A nge l , J .L . ( 1 984 ) . H ea l th a s ac r uc ia l f a c to r i n t h e c hanges f r om h un t ing t o d eve loped f a rm ing i n t h e E a s ter n Med i te r ranean . I n P a l eopa tho logy a t t h e O r ig ins o fA g r icu l tu re . e d s . M .N . C ohen a nd G .J . A rme lago s .p p .5 1 -73 .O r lando : A cadem ic P re s s . B a rke r , G . ( 1 985 ) . P reh i s to r ic f a rm ing i n E u rop e . C ambr idge : C amb r idge U n ive r s i ty P re s s . B ende r , B . ( 1 978 ) . G a the r e r -hun te r t of a rme r : as o c ia l p e r spec t ive . Wo r ld A rchaeo l . 1 0 (2 ) :203 -222 B ende r , B . ( 1 985 ) . Eme rgen t t r iba l f o rma t ion s i n t h e Ame r ican M idcon t inen t . Am .A n t iq .5 0 ( 1) :52 -62 .

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A cknow ledgemen t

Z ve leb i l , M . a nd R ow ley -Conwy , P .( 1 984 ) . T ran s i t ion t o f a rm ing i n n o r the rn E u rop e : ah un te r -ga the re r p e r spec t ive .N orweg ian A rchaeo l .R ev . 1 7 :104 -28 . Z ve leb i l , M . a nd R ow ley -Conwy , P . ( 1 986 ) . F o rage r s a nd f a rme r s i n A t lan t ic E u rope . I n H un te r s i n T rans i t ion : Me so l i th ic S oc ie t ie s o f T empe ra te E u ra s ia a nd T he i r T ran s i t ion t o F a rm ing . e d . M . Z ve leb i l . p p . 6 7 -93 . C amb r idge : C amb r idge U n ive r s i ty P r e s s . Z ve leb i l , M . a nd Z ve leb i l , K .V . ( 1 988 ) . A g r icu l tu ra l t r ans i t ion a nd I ndo -Eu rop ean d i sp e r sa l s . A n t iqu i ty . 6 2 :574 -583 . 2 i v a n o v i c , S . ( 1 975 ) . A n o te o n t h e a n th ropo log ica l c ha rac te r i s t ic s o f t h e P ad ina p opu la t ion . Z e i ts chr if t Mo rpho l .A n thropo l . 6 6 :161 -175 .

C . Me ik le john w ou ld l i ke t ot h ank M . Z ve leb i l f o r t h e i n v i ta t ion t o t h e TAG88 c onf e rence a nd t h e s ugge s t ion t h a t t h i s j o in t p ape r w ou ld b e u s efu l . A t t endance a tt h e S hef f ie ld m e t ing w a s m ade p o s s ib le b y aR e sea rch F u r lough G ran t f r om t h e U n ive r s i ty o f W inn ipeg . A sp ec t s o f t h e w o rk r e po r ted w e re m ade p os s ib le b y g r an t s 4 10 -87 1045 a nd 4 10 -89 -0376 t oC . Me ik le john b y t h e S oc ia l S c ience s a nd H uman i t ie s r e sea rch C ounc i lo fC anada .

1 4 3

T ABLE 1 .1

CAR IES RATES I N THE EUROPEAN MESOL ITH IC R eg ion /S i te

C ar ie sN umber

C ar ie s R a te [ g o ]

S ka teho lm I ,S weden S ka teho lm I ,S weden V edbaek -Bogebakken ,D enma rk O the rn on -ceme t e ry m a te r ia l s T o ta lS cand inav ia

5 / 848 0 / 491 0 / 357 1 / 203 6 / 1899

0 . 59 0 . 0 0 . 0 0 . 49 0 . 315

T dv iec ,F r ance H oed ic ,F rance O the rn on -ceme te ry m a t e r ia l s T o ta lM id -La t i tude E urope

8 / 352 7 / 133 0 / 181 1 5 /666

2 . 27 5 . 26 0 . 00 2 . 25

Mo i ta d oS eba s t iao ,P o r tuga l C abeco d aA r ruda ,P o r tuga l U zzo a nd Mo la ra ,I t a ly E lC ing le V e rme i l ,S pa in O the rn on -ceme te ry m a t e r ia l s T o ta l Med i t erranean E urope

1 0 7 /889 8 5 / 1176 2 8 /223 8 / 9 2 / 128 2 30 /2425

1 2 .04 7 . 23 1 2 .56 8 8 .88 1 . 57 9 . 48

TOTAL EUROPEAN MESOL ITH IC

2 51 14990

5 . 03

S ource sa si n Me ik le john e ta l . ( 1 988 )e xcep t :

S ka teho lm E lC ing le V erm e i l

A lexande r sen , 1 9 88 T u rbon , 1 986

T he se f i gur e st a ke p r eced ence o ver t h o se p ub l i shed i n Me i k l e j ohn , e ta l . (1988 ) .

1 4 4

V edbae l c ,D enma rk

S ka teho lm , S wed en

C V ; • • 4

‘ 0 1 : o t i

1 45

1 1 l e ; 1 (

S ande r rnokha ,R u s s ia

Abo ra ,L a t v ia

F ranch th iC ave ,G reece

O ther b ur ia ls t ie s

Mo i ta d oS ebe s t iäo ,

Med i terran ean ' p a t tern

S i tes c harac t er is t ic o ft he ' Wes t

0 C • O

C eme te r ie sa nd B u r ia l si n Me so l i th ic E u rop e

S i te sc harac t er i s t ic o f' Nor th E as tE uropean 'p a t tern .

•

1 4

• c ;