Sustainable Management of Nematodes in Agriculture, Vol.1: Organic Management (Sustainability in Plant and Crop Protection, 18) 3031099427, 9783031099427

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Sustainability in Plant and Crop Protection 18

Kamal Kishore Chaudhary Mukesh Kumar Meghvansi   Editors

Sustainable Management of Nematodes in Agriculture, Vol.1: Organic Management

Sustainability in Plant and Crop Protection Volume 18

Series Editor Aurelio Ciancio, Sezione di Bari, Consiglio Nazionale delle Ricerche Istituto per la Protezione delle Piante, Bari, Italy

The series describes new perspectives recently developed in pest and disease management, based on innovative tools emerging from basic and applied research. The volumes will aim at interested readers involved in plant protection and crop management, for whom soil biodiversity, crop sustainability and, in general, organic approaches are fundamental. Different cropping systems will be treated by researchers involved in cutting edge studies worldwide. A number of basic issues including sustainability, life-cycle assessment, evolution, plant nutrition and organic cropping technologies will provide a common framework, within which different components of the crop production cycle will be focused on. These will range from roots and endophytes to pest and disease control, through the management of soil microbiome and fertility. These issues will be examined at the field and crop levels, including the effects of invasive species and climate changes on agroecosystems. Recent advancements in massive sequencing will represent the basis of dedicated volumes, dealing with transcriptomics and related approaches. They will illustrate the potentials and benefits of extensive DNA and RNA data analyses and studies, for practical purposes in crop protection, disease management and food production. Contributions on any of the above cited topics are welcome. Potential Editors proposing a new volume are requested to contact the Series Responsible Editor and provide a short CV (400 words max) listing a selection of their most significant publications. In order to broaden the base of contributors and avoid redundancies, only one volume per Editor is allowed. Exceptionally, in case of many contributed chapters, a two-issues volume can eventually be considered.

Kamal Kishore Chaudhary Mukesh Kumar Meghvansi Editors

Sustainable Management of Nematodes in Agriculture, Vol.1: Organic Management

Editors Kamal Kishore Chaudhary Former Faculty, Jaipur National University Jaipur, India

Mukesh Kumar Meghvansi Bioprocess Technology Division Defence R&D Establishment Gwalior, Madhya Pradesh, India

ISSN 2567-9805     ISSN 2567-9821 (electronic) Sustainability in Plant and Crop Protection ISBN 978-3-031-09942-7    ISBN 978-3-031-09943-4 (eBook) https://doi.org/10.1007/978-3-031-09943-4 © The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed. The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. The publisher, the authors, and the editors are safe to assume that the advice and information in this book are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the editors give a warranty, expressed or implied, with respect to the material contained herein or for any errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. This Springer imprint is published by the registered company Springer Nature Switzerland AG The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland

Preface

Plant parasitic nematodes cause considerable losses to agricultural crops, globally. The estimated yield loss due to these tiny pests is more than US$ 150 billion annually. In terms of percentage, this estimated global yield loss is around 10% annually. In order to fulfill the demand of growing population, significant technological efforts are being made to provide sustained agricultural productions that may also include use of synthetic nematicides. Majority of farmers actually use such nematicides to safeguard the food and commercial products they produce. Synthetic nematicides are effective and are known to reduce the nematode population in agro-ecosystem in a short term. However, their extensive and indiscriminate use poses human health and environmental problems. Nematicides exposure is one of the foremost concerns about environmental safety worldwide. Continued exposure either through food products and fodders or through work exposure is currently posing a severe health hazard throughout the world. In the process of making decisions concerning the usage of nematicides, both new and old, risk assessment remains critical. Considering the ill effects of synthetic nematicides and rising concerns about economic and ecological consequences of their use, global research efforts are looking for environment-benevolent and sustainable alternative approaches for nematode control in sustainable agriculture. Organic amendment is one of the promising strategies that can play a significant role in sustainable nematode management, building up at the same time the soil fertility for sustainable agriculture. Various research efforts have highlighted the potential role of different organic amendments. These include green manures, animal manures, composts, plant products, and essential oil, among others, applied in effective nematode management strategies and improving plant growth. In addition, investigation on agricultural wastes, such as dried-crop residues, and industrial by-­ products, such as oil cakes, sawdust, cellulosic waste, and sugar-cane bagasse, has provided encouraging results. These organic amendments can contribute to the suppression of plant parasitic nematodes directly or indirectly, through their effects on

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nematode eggs, juveniles, and adults. They can also inhibit the nematode population by sustainable, although temporary, changes in soil physicochemical properties, including pH and organic matter. It is therefore important to know the influence mechanism of organic amendments on nematode suppression. In spite of the established potential of organic amendments in sustainable nematode management, the scientific information on the topic is scanty. The present volume, Organic Management of Nematodes in Agriculture, therefore, is an endeavor to synthesize the latest, in-depth, and critical information on the organic products available today for nematode management. Here, we provide 16 chapters contributed by international professionals divided into two parts. Part I, with eight chapters, covers paradigms and mechanisms of nematode management through organic means. Part II, comprising of eight chapters, deals with the applied aspects of organic nematodes management and regional case studies/success stories. We believe that the present volume provides a balanced view of basic as well as applied aspects that will be useful to students, researchers, and scholars working in the field of sustainable nematode management. Further, we consider that the data and information herein gathered and discussed will assist policy makers and administrative authorities all over the world, when regulating sustainable nematode management strategies, thereby minimizing use of chemicals-based approaches. The volume editors would like to express their sincere gratitude to all contributors for submitting their work and timely responding to all the post-submission editorial queries. We have received numerous insightful and constructive inputs from researchers all across the world on this subject while editing this book, for which we are sincerely grateful to them. It was indeed a memorable experience reading through the exciting knowledge synthesized by the authors in their chapters. We would also like to thank the editorial as well as production team at Springer, particularly Mr. Prasad Gurunadham (Project Coordinator, Books), Zuzana Bernhart (Executive Editor, Plant Sciences – Books), Mariska Van Der Stigchel, and Albert Paap, for their critical evaluation, encouragement, and constant whole-hearted support. Dr. Kamal K. Chaudhary would like to place on record his deep sense of gratitude to his research mentor late Dr. R. K. Kaul, Principal Scientist, Central Arid Zone Research Institute, Jodhpur, for his guidance and training, which was the motivation to bring this volume as a tribute to the Indian nematologist par excellence. Dr. Chaudhary also wishes to thank his family members, Mrs. Kesher (mother), Mrs. Manju Chaudhary (wife), Mr. Pulkit Chaudhary (son), and Miss Jyoti Chaudhary (daughter), for their unconditional love, patience, encouragement, and incredible support during this period. Last but not the least, Dr. Chaudhary also wishes to thank all friends, staff members, and relatives for their encouragement during the entire editing tenure. Dr. Mukesh K. Meghvansi wishes to thank Mrs. Manju Meghvansi (wife), and Miss Lakshita Meghvansi and Parnika Meghvansi (daughters) for their love, patience, understanding, and moral support.

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Last but not the least, the volume editors wish to thank Dr. Aurelio Ciancio, Series Editor – Sustainability in Plant and Crop Protection, for his critical evaluation, constant support, and encouragement. Jaipur, India

Kamal Kishore Chaudhary

Gwalior, Madhya Pradesh, India

Mukesh Kumar Meghvansi

Contents

Part I Organic Management of Nematodes: Paradigms and Mechanisms 1

Use of Natural and Residual Resources for the Sustainable Management of Phytonematodes: Challenges and Future Trends����������������������������������������������������������������    3 Thales Lima Rocha, Vera Lucia Perussi Polez, Lívia Cristina de Souza Viol, Reinaldo Rodrigues Pimentel, Danielle Biscaia, and Jadir Borges Pinheiro

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Organic Nematicides: A Green Technique and Its Overview for Nematode Pest Management������������������������������   39 Faryad Khan, Mohammad Shariq, Mohd Asif, Taruba Ansari, Saba Fatima, Arshad Khan, Mohd Ikram, and Mansoor Ahmad Siddiqui

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 Nematode Management Prospects in Composting ������������������������������   67 Fisayo Yemisi Daramola, Samuel B. Orisajo, and Osarenkhoe Omorefosa Osemwegie

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Biochemical/Molecular Mechanisms Associated with Nematode Management Through Organic Amendments: A Critical Review������������������������������������������������������������   87 John Fosu-Nyarko, Rhys G. R. Copeland, Sadia Iqbal, and Michael G. K. Jones

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Agroindustrial By Products Suppressing Plant-Parasitic Nematodes����������������������������������������������������������������������  117 Alixelhe Pacheco Damascena, Marylia Gabriella Silva Costa, Júlio César Antunes Ferreira, and Silvia Renata Siciliano Wilcken

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 Nematode Management by Humic Acids����������������������������������������������  135 Seenivasan Nagachandrabose

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Conventional and Organic Management as Divergent Drivers for Plant Parasitic Nematodes Control������������������������������������  157 Kanika Khanna, Vandana Gautam, Dhriti Kapoor, Nandni Sharma, Pooja Sharma, Tamanna Bhardwaj, Puja Ohri, and Renu Bhardwaj

Part II Organic Management of Nematodes: Global Case studies and Success Stories 8

Plant Extracts and Their Effects on Plant-­Parasitic Nematodes, with Case Studies from Africa ������������������������������������������  189 Ebrahim Shokoohi

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Non-chemical Management of the Citrus Nematode, Tylenchulus semipenetrans (Nematoda: Tylenchulidae) ����������������������  217 Reza Ghaderi and Manouchehr Hosseinvand

10 Organic  Management of Rice Root-Knot Nematode, Meloidogyne graminicola ������������������������������������������������������������������������  247 Ziaul Haque and Mujeebur Rahman Khan 11 Strategies  for the Organic Management of Root-Knot Nematodes (Meloidogyne spp.) in Vineyards Under Desert Conditions in the North Coast of Peru��������������������������������������  269 César Augusto Murguía Reyes 12 Organic  Management Strategies for Nematode Control in Florida Plasticulture��������������������������������������������������������������  293 Johan Desaeger, Kaydene Williams, and Erin Rosskopf 13 Eco-friendly  Management of False Root-­Knot Nematode Nacobbus aberrans: An Overview����������������������������������������  327 Edgar Villar-Luna, Olga Gómez-Rodríguez, Hernán Villar-Luna, Liliana Aguilar-Marcelino, Laith Khalil Tawfeeq Al-Ani, and Ernesto Fernández-Herrera 14 Organic  Amendments and Other Strategies for Management of Meloidogyne spp. and Nacobbus aberrans in Horticultural and Orchard Crops: The Mexican Experience ������������������������������������  343 Ignacio Cid del Prado-Vera, Marco Antonio Magallanes-Tapia, Raúl Velasco-­Azorsa, and Arely Pérez-Espíndola

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15 N  on-conventional Management of Plant-­Parasitic Nematodes in Musaceas Crops ��������������������������������������������������������������  381 Donald Riascos-Ortiz, Ana T. Mosquera-Espinosa, Francia Varón de Agudelo, Claudio Marcelo Gonçalves Oliveira, and Jaime Eduardo Muñoz Flórez 16 Neem  Cake Amendment and Soil Nematode Spatio-Temporal Dynamics: A Case Study in the Brazilian Semiarid Region�����������������������������������������������������������  423 Diego Arruda Huggins de Sá Leitão, Ana Karina dos Santos Oliveira, Douglas Barbosa Castro, and Elvira Maria Régis Pedrosa

Part I

Organic Management of Nematodes: Paradigms and Mechanisms

Chapter 1

Use of Natural and Residual Resources for the Sustainable Management of Phytonematodes: Challenges and Future Trends Thales Lima Rocha , Vera Lucia Perussi Polez , Lívia Cristina de Souza Viol , Reinaldo Rodrigues Pimentel , Danielle Biscaia , and Jadir Borges Pinheiro

Abstract  The growing demand for safe food associated with increased restrictions for the use of synthetic agrochemicals in different cultures has led to the development of more sustainable technologies for control of pests, such as phytonematodes. In this sense, natural products and residues are rich sources of compounds and nutrients that can contribute to productivity and the control of these phytoparasites. Thus, this chapter presents the scientific bases, examples of success, mechanisms of action, advantages and disadvantages regarding the use of: (i) botanical and fungal resources; (ii) management with cover crop and industrial plant residues; (iii) resources from animals and agro-industrial wastes; and (iv) blends. Additionally, a discussion concerning natural or recycled products is proposed, indicating the challenges and trends. In this context, challenges concern: (i) biodiversity conservation, (ii) quality system (e.g. rules and standardization) to guarantee reproducibility, repeatability, reliability, stability, efficiency, and safety, (iii) government policies, (iv) market regulations, public and private institutions integration. Finally, we discuss trends regarding nanotechnology-based green chemistry,, the use of blenders, the Integrated as well as Holistic Pest Management. These trends together integrate

T. L. Rocha (*) · V. L. P. Polez · L. C. de Souza Viol Embrapa Genetic Resources and Biotechnology, Parque Estação Biológica, Brasília, Brazil e-mail: [email protected]; [email protected] R. R. Pimentel Faculty of Agronomy and Veterinary Medicine (FAV) Ala Central, University of Brasília, Brasília, Brazil D. Biscaia · J. B. Pinheiro Embrapa Vegetables, Brasília, Brazil e-mail: [email protected]; [email protected] © The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 K. K. Chaudhary, M. K. Meghvansi (eds.), Sustainable Management of Nematodes in Agriculture, Vol.1: Organic Management, Sustainability in Plant and Crop Protection 18, https://doi.org/10.1007/978-3-031-09943-4_1

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the farmer in designing solutions for pest control, minimizing socioeconomic and environmental impacts, and customer satisfaction. Keywords  Phytonematodes · Control · Biopesticides · Plant extracts · Essential oil · Fungi extracts · Cover-crop plants · Agro-industrial waste · Blends

1.1 Introduction World agriculture provides food for billion human beings. In this sense, population growth has required strategies that allow an increase in food productivity to ensure global food availability (The State of the World’s Biodiversity for Food and Agriculture, 2019). However, one of the central challenges in food production is pest control. Among them, plant-parasitic nematodes such as root-knot nematodes (Meloidogyne spp.), cyst nematodes (Heterodera spp. and Globodera spp.), and lesion nematodes (Pratylenchus spp.) represent one of the main causes of annual crop losses (Jones et al., 2013). These phytoparasites (Fig. 1.1) affect several crops with high added value and, therefore, cause global annual losses on the order of million dollars (Mesa-Valle et al., 2020). Currently, the main strategy of nematode control is the use of synthetic nematicides (Lengai et al., 2020). However, most of these agrochemicals are potentially toxic to both human and animal health and the environment. For this reason, the use of many nematocides has been restricted in several countries (EC, 2013; USEPA, 2009). To overcome this limitation, and also to ensure the availability of safer food supplies, several strategies have been developed. Among them, the use of plant and fungus resources, animal and plant agro-industrial residues, the Integrated Pest Management (IPM), and blends provided some options with a great potential for applicability. In a “green” chemistry èersèective, the use of sustainable technologies with biocidal action aimed at reducing or minimizing the adverse impacts of toxic products on farmers and consumers health, as well as the environment. In this sense, the quality and safety of agricultural products are a worldwide trend. It is worth mentioning that, throughout this process, the social awareness for consumption of safe and environmentally low impact food has been crucial. This social movement has contributed in the last decades to major changes in the industry and market of conventional pesticides. Global biodiversity and its ecosystems constitute a very rich source of biologically active materials which could be used in traditional crop protection (The State of the World’s Biodiversity for Food and Agriculture, 2019). In this context, Brazil biodiversity resourcescan generate global proactivity in the areas of science, technology, and innovation through a number of biological inputs for sustainable agriculture. It offers a great opportunity to explore new molecules, to find out possible distinct functions as well applications. These varieties of molecules can be obtained

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Fig. 1.1  Soybean field exhibiting plants with stunting and yellowing, or symptoms of chlorosis due to infestation by soybean cyst nematodea (Heterodera glycines) (a); Nematode resistance structure: cyst of Heterodera glycines (b); Symptoms in potato tubers due to infestation by a root-­ lesion nematode (Pratylenchus spp.) (c); Symptoms in carrot roots due to the presence of root-knot nematodes (Meloidogyne spp.) (d); Melon roots showing galls (e) due to parasitism by a Meloidogynes sp. Symptoms in white sweet potato roots caused by Meloidogyne sp. (f). (Authors: Danielle Biscaia and Jadir Borges Pinheiro)

by prospecting different biomes and/or germplasm or microorganism banks. In addition to these sources, animal and agro-industrial wastes also represent another important source of potentially useful molecules. Thus, natural resources and residues are important sources of complex molecules, with a high capacity to interact with different biological systems to control pests such as phytonematodes, using strategies and rules of “green” chemistry.

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Formulations are essential to obtain more sustainable and multifunctional technologies. In this sense, nanotechnology-based tools  – i.e. nanocarriers such as metallic nanoparticles, polymeric nanoparticles, nanoemulsions, among others  – can carry compounds, extracts, or blenders resulting in a more controlled release and in a lesser amount of material to be used for phytonematodes control. In this context, the establishment of a quality system is fundamental to obtain natural sources, residues, or blends and their formulations with reproducibility, repeatability, reliability, stability, efficiency resulting in safe food and customer satisfaction. Thus, this strategy will follow quality regulatory mechanisms that aim to increasingly meet the demands of society, the media, and national/international trade, which currently calls for safe food products with a environmental impact. Another powerful strategy is correlated to Integrated Pest Management (IPM) or Holistic Pest Management (HPM) that can integrate, during the crop development, strategies such as nanotechnology and blends, resulting in a more efficient pest control, also guaranteeing human health and environment protection.

1.2 Plant Extracts Plant biodiversity represents an ample source of biologically active materials that could be exploited in crop protection vs a wide range of pests and pathogens, including phytonematodes (Caboni & Ntalli, 2014; Sivasubramaniam et al., 2020; Tiku, 2020). These active materials, denominated extracts, are obtained from plant parts such as barks, leaves, roots, flowers, fruits, seeds, cloves, rhizomes, stems, and others. Plant extracts derived from distinct botanical families have been used as a basis for the development of biopesticides. In this sense, Brazil 6 distinct Biomes (Amazon, Atlantic forest, High Savannah, Semi-arid, Prairie and Wetlands) represent a vast reservoir of plants and, ultimately, extracts and compounds, with a repertoire of distinct biopesticides (Guerra et al., 2020). When compared with conventional synthetic pesticides, biopesticides are normally less toxic. They usually affect only the targeted pest and closely related organisms, are often effective in relatively small amounts and decompose faster, resulting in a reduced exposure. In general, plant extracts demonstrate a significant risk reduction among pest species as concerns the development of resistance mechanisms, due to a reduced selection pressure. In addition, a remarkable reduction of pest management costs has also been observed, induced by lower infestation pressure, lower accumulated residues and damage, with improved crop protection and yields. Moreover, higher crop protection standards can be achieved based on the application of plant extracts resulting from the positive interactions between botanical products and beneficial macro and microorganisms, in greenhouse and field crops. A lesser toxicological and ecotoxicological risk is also observed for field workers, consumers, wildlife, and the environment. Plant extracts also allow a permanent establishment of beneficial microorganisms, in greenhouse as in field crops.

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Finally, they promote an agro-ecosystem that stimulates the establishment and growth of beneficial microorganisms, associated with the crop and soil environment. On the other hand, some limitations may be related to the use of botanical pesticides. In this sense, the quality of the raw material used for production is crucial to obtain promising results, as the plant material used for production should be selected at the proper time (seasonality). Moreover, the active ingredients of the various plant parts exhibit a huge variability in different geographic areas, limiting the product standardization. Another concern is related to the fact that the majority of botanical pesticides display fast degradation and consequently demand higher application rates and frequency. In addition, regulatory authorization for botanical extracts is expensive and time-consuming, and it is still based, mainly, on the chemical pesticide approval model. The limitation in the production of plant material with nematotoxic effects on a large scale represents another critical issue. Poor economic feasibility for many products obtained from complex and expensive processes, especially when the effective molecules have low solubility, exemplifies a further drawback. Various plant extracts and toxic substances have been evaluated for phytonematodes control (Oka, 2010). Among them, extracts from plants belonging to the following botanical families stand out: Brassicaceae (Brassica carinata and B. napa), Fabaceae (velvet bean, Mucuna spp. and sunn hemp, Crotalaria juncea), Asteraceae (marigold, Tagetes spp.), and Euphorbiaceae (castor bean, Ricinus communis) (Wang et al., 2002; Hooks et al., 2010; Mokrini et al., 2010; Umar et al., 2010). The common bioactive compounds in botanical pesticides are mostly secondary metabolites, such as steroids, alkaloids, tannins, terpenes, phenols, and flavonoids among others. In vitro bioassay with aqueous leaf extracts from Bael (Aegle marmelos) and Neem (Azadirachta indica), at a concentration of 88.53% and 80.31%, exhibited mortality toward the rice root-knot nematode M. graminicola, ranging from 80 to 88%, after 48 h exposure (Dongre & Sobita, 2013). In greenhouse experiments, the extracts from Bael and Neem also exhibited a high reduction of root gall (35.33 and 22.66) induced by M. graminicola in rice, when compared to the control treatment (nematode alone). Another study conducted by Ntalli et  al. (2020b) in vitro showed significant nematicidal activity of aqueous extracts from leaves and wooden stems of Stevia rebaudiana (fam. Asteraceae) vs M. incognita and M. javanica. In addition, in vivo bioassays conducted in greenhouse conditions showed substantial efficacy of the leaf powder (95% at 1 g/kg) followed by stems. Rocha et  al. (2017) reported that the aqueous extract (1  mg/mL), the external dialysate (0.5  mg/mL) as well as some compounds (0.01  mg/mL) isolated from Canavalia ensiformis seeds exhibited, in in vitro conditions, a nematicidal >85% mortality effect on M. incognita. It is worth mentioning that the external dialysate showed thermostability, low toxicity against bovine red blood cells, and did not affect non-targeted organisms at the same concentration that killed the nematodes. Under greenhouse conditions, the external dialysate decreased the M. incognita egg masses by 82.5% in tomato plants.

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According to Ismail et al. (2020), evaluation through in vitro bioassay of nematicidal effects, using the aqueous extracts of Allium sativum, Urtica dioica, Sophora mollis, Ephedra intermedia, and Tanacetum baltistanicum against M. incognita, revealed a mortality of 75–95% at concentrations of 0.125–1.0%, after 72 h exposure. Furthermore, methanolic extracts from the aerial part of Datura stramonium, and seeds of Solanum nigrum, showed nematicidal activity against M. incognita and M. javanica (Oplos et al., 2018). Some compounds from plant extracts have also shown effects against phytonematodes. Studies conducted by Aoudia et al. (2012) using the phenolic compounds p-coumaric acid and p-hydroxybenzoic acid, present in the aqueous extract of the Melia azedarach fruit pulp, showed in vitro a nematicidal effect (EC50/48h = 840 and 871 μg/mL) against M. incognita. The effect of 49 phenolic compounds on M. incognita, under in vitro and greenhouse conditions, has been evaluated by Oliveira et  al. (2019). D-(-)-4-­ hydroxyphenylglycine, t-butylhydroquinone, L-3-(3,4)-dihydroxyphenylalanine, sesamol, 2,4-dihydroxyacetophenone and p-anisaldehyde were the most effective with (LC50: 365, 352, 251, 218, 210, and 85  μg/mL, respectively) under in vitro conditions. Additionally, it was also shown that hydroquinone (at 3.5  mg/plant) reduced M. incognita populations and galls by up to 99%, levels similar to the nematicide Carbofuran (at 1.2 mg/plant). Plant extracts can be used as sources for the “green synthesis” of nanoparticles to control phytopathogens (Kalaiselvi et al., 2017; Abbassy et al., 2017; Silva & Bonatto, 2019; Hernández-Díaz et al., 2020). Green synthesis uses relatively non-­ toxic, biodegradable, and low-cost chemicals as primary sources to synthesize nanomaterials. Biological resources (plants, microorganisms, various agricultural by-products, among others) can be used from a biological organism or part of it (organs, tissues, cells, biomolecules or, metabolites) as potential sources for the green synthesis of metallic nanoparticles (Sharma et al., 2009; Silva et al., 2015; Lee et al., 2020). Some eco-friendly nanomaterials have potential to control phytonematodes. Aqueous extracts from two tropical plants, Curcuma longa (fresh tubers) and A. indica (fresh leaves), were used for the green synthesis of silver nanoparticles (AgNPs) that showed dose-dependent toxicity against M. incognita (Kalaiselvi et al., 2017). The lethal concentration (LC50) after 72 h of AgNPs for A. indica was 6.22 mg/L, and for C. longa at 0.54 mg/L. Comparison of these two green synthesized AgNPs, showed a higher mortality rate of the C. longa extract synthesized AgNPs (Kalaiselvi et al., 2017). Another example are the extracts of Conyza dioscoridis, that showed nanoformulations more toxic than the crude extract vs M. incognita (Abbassy et al., 2017). Botanical pesticides contain bioactive compounds that exhibit different modes of action against phytonematodes. These modes of action, related to the majority of plant-derived bioactive compounds, are complex and little known. The extracts can act as attractants and repellents, or as nematicidal/inhibitory agents, defense elicitors, hatching stimulants/inhibitors, agents of proteins denaturation, and other effects, depending on the type of botanical compound and pest (Sikder & Vestergård, 2020). In this context, nematicidal properties, such as the inhibition of egg hatching

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and the suppression of nematode populations, are commonly found in botanical pesticides. Some of them can also affect the population of other microorganisms in soil, which further affect the survival of the nematode eggs and juveniles (Khan et al., 2008). Moreover, some compounds can also kill second-stage juveniles (J2), reduce egg masses and galling and/or affect the phytonematode population in its ecosystem (Kepenekci et al., 2016). For instance, plant extracts obtained from B. napus, L. camara, T. erecta, and A. indica can inhibit eggs hatching of M. incognita, leading to immobilization and ultimately J2 death. The complex mechanism related to inhibition of root-knot nematodes egg hatching, motility and J2 mortality may be associated with the presence of plant alkaloids, tannins, and glycosides (Akyazi, 2014). Moreover, a considerable number of phytochemicals found in botanical extracts exhibit a lipophilic property allowing them to straightforwardly dissolve the nematodes cytoplasmic membrane, thus interfering with protein structures accountable for nematode growth, development and survival (Pavaraj et al., 2012). Paradoxically, despite the massive amount of research and scientific publications demonstrating the effectiveness of botanical extracts, fractions and compounds in phytonematodes control, they have not been translated yet into commercial products. So far, only a few products exhibiting nematicidal activity based on plant extracts have been made available on the world market. They are based on neem (A. indica) and garlic extracts (Allium sativum). The reasons concerning this problem are related to severe barriers that include: (i) difficulties in scaling-up production of plant materials; (ii) standardization of extracts exhibiting high chemical complexity; (iii) regulatory barriers to commercialization; (iv) slow action of various botanical materials; (v) limited residual action; (vi) accessibility of competing products (Khater, 2012). In addition to this scenario, it is noteworthy to mention that there are still a considerable number of plant species not yet explored around the world, especially in the Brazilian biomes.

1.3 Essential Oil The essential oil (EO) is the product obtained from a vegetable raw material, usually by steam distillation or mechanical extraction processes. EOs are formed by a mixture of volatile compounds responsible for the characteristic aroma of certain plants, found in glandular brushes, papillae, or in secretory cells and intercellular channels, or present as fluid droplets in leaves, stems, bark, flowers, roots and/or fruits. These lipophilic components commonly found in EOs include terpenes, sesquiterpenes, phenolic compounds, ketones, acids and esters, products of plant secondary metabolism (Figueiredo et al., 2008; Koul et al., 2008; Butnariu & Sarac, 2018). Several studies indicate that the terpenes and phenolic compounds present in EOs are the mostly responsible for plants defense from phytopathogens (Bakkali et al., 2008). In general, EOs are not toxic to mammals, birds, fish at low concentrations and are not harmful to the environment. Formulations based on EOs are one of the best

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alternatives to the use of synthetic chemicals for nematode control, especially those that use emulsions or encapsulation, because of prolonged and improved nematicidal efficacy at the field level. In these conditions the EOs are protected from rapid degradation and evaporation from soil, thus allowing a prolonged effect on nematodes (Figueiredo et al., 2008; Laquale et al., 2018). In addition, it is worth mentioning the expectation of using these products in small areas, in organic productions and mainly in the cultivation of vegetables, where many of the species produced have extremely short planting-to-harvest cycles, and are consumed “in natura”, thus reducing the risk of chemical residues in harvested products. The commercial use of products based on EOs requires the availability of sufficient amounts of plant material for continued production and the standardization of final products. The EOs composition may in fact present great qualitative and quantitative variations, in relation to different agronomic factors, plant species, and extraction techniques (Laquale et al., 2015). Furthermore, the high volatility of EOs may result in a low permanence time in soil, and an ineffective action in field conditions. The use of emulsions or encapsulations in formulations aim at reducing or solving these challenges, improving solubility and promoting the EO controlled release. Thus, products used outdoors may require frequent re-applications or adequate protection and release strategies in field situations, which can increase production costs (Koul et al., 2008; Domingues & Santos, 2019). Research related to the EO effects on phytonematodes has shown promising results. EOs from Mentha spp., Eucalyptus spp., Cymbopogon spp., Pelargonium graveolens, and Ocimum basilicum demonstrated high toxicity to root-knot nematodes. Jardim et al. (2020) tested the effect of EO from garlic (A. sativum) obtained by hydrodistillation in the control of M. incognita in vitro and in greenhouse. This EO was more active at 63 μg/mL against M. incognita eggs and J2 than the nematicide Carbofuran at 173  μg/mL.  Infectivity and reproduction of M. incognita in tomato plants cultivated in substrate inoculated with nematode eggs and treated with 0.2 mL/L of substrate were statistically equal to those observed with 0.25 g of dazomet/L of substrate. These results confirm the activity of the garlic EO and its components against M. incognita, suggesting its potential as a new fumigant nematicide. Kalaiselvi et al. (2019) evaluated in vitro and greenhouse the nematicidal activity against M. incognita of the EOs obtained from Artemisia nilagirica grown at different altitudes. EOs obtained from high and low altitudes showed a significant difference in the nematicidal activity, with a lethal concentration (LC50/48 h) of 5.75 and 10.23 μg/mL, respectively. The EOs of A. nilagirica reduced the infection of tomato roots by M. incognita and significantly promoted the growth of plants in the greenhouse. EOs from Monarda didyma and M. fistulosa and their main compounds: carvacrol, γ-terpinene, o-cymene, and thymol, were evaluated in vitro and greenhouse for their activity against M. incognita and Pratylenchus vulnus (Laquale et al., 2018). Both EOs were strongly active against J2 of M. incognita, as only 1.0 μL/mL of LC50 was obtained after a 24 h exposure to both EOs, while a lower activity was

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recorded in P. vulnus (15.7 and 12.5 μL/mL of LC50 for M. didyma and M. fistulosa, respectively). The almost similar chemical composition of the two tested EOs suggests that their different behavior to the two nematode species could not be only attributed to EOs chemical composition, as variability intrinsic to the nematode species should be also considered (Laquale et al., 2018). Laquale et al. (2015) evaluated the EOs of Eucalyptus citriodora, E. globulus, Mentha piperita, Pelargonium asperum, and Ruta graveolens against M. incognita on potted tomato in the greenhouse, at a concentration of 50, 100, and 200 μL/kg soil for each EO. At 50 μL/kg, the number of galls and M. incognita eggs on tomato roots were significantly lower than the non-treated control, but only in soil fumigated with the EOs from E. globulus and P. asperum. The EOs mechanism of action against nematodes is still unclear and it is likely correlated with the chemical structure of its components (Laquale et al., 2015). The presence of volatile monoterpenes and other lipophilic phytochemicals provides an important defense strategy against nematodes. Some EOs lipophilic phytochemicals can easily dissolve the nematode cytoplasmic membrane and directly interfere in the synthesis of proteins underpinning its growth, development, and survival (Lengai et al., 2020). Different hypotheses have been proposed for the mechanism of action on nematodes in the literature, including a neurotoxic mode of action, disruption and change of permeability of nematode cell membranes or the inhibition of AChE (acetylcholinesterase) activity. Further studies are needed to clarify this question (Kostyukovsky et al., 2002; Andrés et al., 2012; Laquale et al., 2018; Saroj et al., 2020; Oka et al., 2000). Despite the increase in research work and scientific publications on EOs nematicidal properties, few studies have been conducted in the field, and nematicidal products based on EOs are not available on the world market. This fact may be due to difficulties in standardizing products and field applications, regulatory barriers to commercialization, as well as production costs higher than common marketed products available. Given the global context of sustainable agriculture, the prospect of using commercial nematicides based on EO must overcome these challenges, with more in-­ depth research in the field, dosage studies, feasibility, use of new application technologies and product standardization. In addition, given Brazil extensive biodiversity, few species of native plants have yet been the subject of research involving EOs in this country.

1.4 Fungal Extracts Fungi are very promising sources of natural compounds for pest control, for promotion of plant growth, and/or to induce disease resistance (Bills & Gloer, 2016; Bogner et al., 2017; Masi et al., 2018; Keswani et al., 2019; Vinale & Sivasithamparam, 2020). Biopesticides can be obtained from living fungi, their extracts (e.g. exudates,

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mycelium) as well as their isolated volatile and non-volatile compounds (Degenkolb & Vilcinskas, 2016; de Souza et al., 2018). Fungi and fungal products are used by several companies for the large-scale production of metabolites, proteins, enzymes, and other compounds. The metabolite-­ producing fungi with biocidal action can be used as bio-factories and have several advantages such as (i) rapid growth (factor dependent on selected species/lineages); (ii) production of more homogeneous chemical compounds (controlled culture conditions); (iii) production in bioreactors (scaling); and iv) contribution to the bioeconomy (use of agro-industrial residues for the cultivation of some groups of fungi as ascomycetes and basidiomycetes) (Lin & Sung, 2006; Silva et  al., 2016). However, the cultivation of fungi has some disadvantages such as: (i) genetic stability maintenance; (ii) homogeneous biological material maintenance; and (iii) contamination during production. Thus, some strategies can be used to solve or significantly minimize these challenges, including (i) methodologies for the fungi storage and tests to verify the genetic stability quality; (ii) protocols for the growth/ development of the fungus (nutrients/substrates, pH, temperature, luminosity, among others); and (iii) processes using aseptic conditions. Metabolite compositions can be altered by some factors such as: (i) species or lineages selection; (ii) biological source selection (e.g. exudates, mycelia) and growth phase; (iii) cultivation conditions (type of substrate/nutrients, pH, temperature, among others); (iv) abiotic (temperature, light, water stress, among others) or biotic stressors (fungi, bacteria); and (v) extraction process and solvents used (Calvo et al., 2002; VanderMolen et al., 2013; Jiaojiao et al., 2018; Khan et al., 2020). Most fungal extracts or metabolites are obtained using reagents such as ethyl acetate and methanol. In this case, these toxic reagents are normally used in both the extraction process and the solubilization step. Strategies to minimize or resolve this negative impact could be (i) to solubilize the extracts obtained in formulations used in sustainable agriculture; and (ii) rely on less toxic or aqueous reagents in the extraction process. In addition, an important point is the characterization of the physical-­ compound properties (solubility, stability, among others) of extracts or compounds that can lead to formulations that do not impact the environment, as well as animals and humans health. In addition, these formulations also aim to maintain the natural product (extract, or compounds) stability, if subjected to highly variable environmental conditions (rainfall, light intensity, pH, presence of other organisms) allowing its use in different regions, or period of the year. Theuse of fungi to obtain metabolites is thus a promising strategy for the production of natural compounds under conditions of scalability and reproducibility, for pest control. In this sense, fungi are sources of metabolites active against nematodes, such as terpenoids, alkaloids, peptides, and aliphatic compounds among others (Li et al., 2007; Nisa et al., 2015; Bogner et al., 2017). Trichoderma species present metabolites with several biocidal applications such as glisoprenin, gliotoxin, gliovirin, viridian, hepteledic acid, polyketides, harzialactones, trichoderm amides and derivatives of α-amino acids. Fungal filtrates of 329 Trichoderma strains showed that 15 presented nematicidal activity against Panagrellus redivivus, and 14 strains exerted control over Caenorhabditis elegans

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(Yang et  al., 2010). The trichodermin compound was identified in a strain (YMF1.02647) that significantly controlled both species (Yang et al., 2010). Volatile organic compound (6-pentyl-2H-pyran-2-one) of Trichoderma sp. (YMF 1.00416) showed nematicidal activity against P. redivivus, C. elegans, and Bursaphelenchus xylophilus (Yang et al., 2012). The extract from Trichoderma viridae (200 mg/mL) showed a significant control against both juveniles and egg-hatching inhibition of M. incognita (Khan et  al., 2020). In this case, the activity was dependent on the growth media as a greater control was observed in the samples obtained from liquid and solid wheat media. Some metabolites can be produced both by plants and some fungi, such as gibberellins and diterpenoid plant hormones (Bömke & Tudzynski, 2009; Bills & Gloer, 2016). The endophytic fungus Fusarium oxysporum 162 showed a promising control activity against M. incognita (Hallmann & Sikora, 1996; Bogner et  al., 2017). Thus, 11 compounds isolated from F. oxysporum 162 extract were analyzed against the root-knot nematode (Bogner et  al., 2017). The compound 4-­hydroxybenzoic showed the best activity (LC50 104 μg/mL) followed by indole-­3-­ acetic acid (IAA) (LC50 117  μg/mL), and gibepyrone D (LC50 134  μg/mL). The positive controls carbofuran and aldicarb showed LC50 72 h values 64 and 180 μg/ mL, respectively. However, the activity of 4-hydroxybenzoic acid was stronger than that of aldicarb. The authors suggested that this fungus can induce resistance against the nematode through two mechanisms: (i) indirect, through increased plant defense by the endophyte-produced 4-hydroxybenzoic acid and indole-3-acetic acid (IAA); and (ii) direct, through the phytohormone (IAA) and the salicylic acid isomer (hydroxybenzoic acid isomers) that could have a dual function, and in the presence of other metabolites produced by the fungus result in the death of M. incognita (Bogner et al., 2017). Basidiomycetes (commonly called mushrooms) are also sources of metabolites with antifungal, antibacterial, antiviral, anti-larvae (larvicidal for mosquito), and nematicidal activities (Sivanandhan et al., 2017). The genus Pleurotus is known to have promising species for nematode control (Sivanandhan et  al., 2017; Sufiate et al., 2017). An example is observed using P. eryngii, as the fungus and its extract reduced the number of intact Panagrellus sp. larvae by 60% and 90%, after 24 h treatment, respectively (Sufiate et al., 2017). The authors, however, did not relate this effect to an enzymatic activity, rather to the presence of other metabolites. In this same work, the extract of P. eryngii reduced the number of intact eggs of M. javanica eggs by approximately 53%, likely through an enzymatic action. Chitinases and proteases activity can affect the structure and development of eggs, causing eggshell ruptures, early hatching, and vacuole formation in eggs and juveniles (Khan et al., 2004; Sufiate et al., 2017). Fungus extracts can be used for the green synthesis of nanoparticles (Silva et al., 2015; Molnár et al., 2018; Barbosa et al., 2019). Aqueous extracts from Duddingtonia flagrans were used as a source for the synthesis of silver nanoparticles (Barbosa et al., 2019). The AgNPs showed action against the larvae of Ancylostoma caninum (parasitic nematode infections) indicating a promising action of fungal extracts as sources for synthesis of AgNPs with nematicidal effects.

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Obtaining commercial products based on fungi extracts is an opportunity to be explored as these organisms can be used as biofactories for the production of homogeneous compounds as well as for production up-scaling. Another important control strategy is the use of fungi associated with extracts or other types of biological materials. In this case, the blends can be formulated using technology strategies such as hydrogels or nanoformulations, for the controlled release and sustainment of natural nematocides. Brazil has a significant fungi diversity that can be used as an important sourcing strategy for phytonematodes control.

1.5 Sustainable Management: Use of Cover Crops and Plant Residues In Brazil, cover plants, as well as organic and industrial residues, have been used in the management of different nematode species, the control of which is of economic importance. It is worth to note that their efficiency in nematodes management depends on several factors such as (a) the size of the area; (b) type of culture; (c) species of cover or antagonistic plants; (d) nematode species and population densities; (e) soil type and degradation; and (f) integrated application of other management practices. In this sense, the demand for this type of nematode management by farmers is more and more focused on advantages provided in management (for example greater nematode tolerance of plants, increased vigor and higher yields) as well as costs reduction in sustainable productions. Cover crops and plant residues are being increasingly used in agricultural systems. Uses have several advantages: recycling nutrients and energy, as well as improving soil physical-chemical conditions for plant growth, and development of microorganisms. Additional benefits of cover crops include nutrients sequesteration (especially nitrates) avoiding their leaching below the crop root zone, suppressing weeds, breaking pest cycles, and providing habitats for beneficial insects and other organisms. Besides these alternatives, cover crops have also been shown to suppress several plant pests including phytonematodes. On the other hand, cover crops used in phytonematodes control have some disadvantages, such as occupying field crop areas, additional planting costs, agronomical limitations, difficult management, functioning as a weed after the cultivation, and multiplying non-targeted phytopathogens. Concerning the use of industrial residues, plants also can be limited by seasonality, as concerns the production of biomasses and the difficulty of large-scale production. Some antagonistic plants have been used successfully as cover crops in the control of phytonematodes including Crotalaria spp. (C. spectabilis), C. juncea L., marigold (T. patula, T. minuta, T. erecta) and mucuna, M. aterrima (Wang et al., 2007; Claudius-Cole et al., 2015). It is worth mentioning that the black mucuna, M. aterrima has proven efficacy vs M. incognita, but does not affect M. javanica (Miamoto et al., 2016). For the control of Pratylenchus spp., the options are more

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limited. In this case, only C. spectabilis, C. ochroleuca, and marigold are indicated (Pudasaini et al., 2006; Cruz et al., 2020). In general, the antagonist plants allow the nematode invasion but prevent their development to maturity. Histopathological analysis of C. spectabilis and C. juncea roots, infected by M. javanica, demonstrated that the nematode induced formation of giant cells in both hosts. However, it was observed that the giant cells were smaller and in fewer numbers when compared to tomato roots, showing that these species are less efficient in meeting the nutritional needs of root-knot nematodes, as compared to susceptible tomato (Silva et al., 1990). Another factor to be taken into consideration is that Crotalaria spp. produces toxic substances, such as monocrotaline, which inhibit the J2 motility in soil. Moreover, the green mass resulting from this plant must be incorporated into the soil, up to approximately 80 days after sowing, before flowering. This procedure is important in avoiding seed production, as well as the high-volume formation of fibrous materials, thus reducing herbicide expenses. Another efficient strategy is the use of cover crops with non-host plants, that keeps the soil moist for a longer time during the autumn-through-winter period. As a consequence, infective stages of phytonematodes remain active. However, because they do not find roots of susceptible plants to inhabit, they end up consuming their own nutritional reserves and dying. Among these types of cover crops the Urochloa species (syn. Brachiaria sp.), U. brizantha, U. decumbens, U. ruziziensis, U. humidicola and U. dictyoneura proved to be non-hosts for M. javanica (Brito & Ferraz, 1987; Inomoto et al., 2007) promoting a considerable reduction in the number of galls per root in tomato plants, that were thus successively cultivated. Dias-Arieira et al. (2003) also demonstrated that U. brizantha and U. decumbens, in addition to different cultivars of Panicum maximum (‘Colonião’, ‘Tanzânia’ and ‘Vencedor’), are potential grasses for rotation in areas infested with Meloidogyne spp. Moreover, the intercropping of Stylosanthes guianensis with yams attacked by M. incognita reduced tuber damage by about 58.3% (Claudius-Cole et al., 2014). Currently, many types of grass have been reported as hosts of P. brachyurus, with a negative impact on the management of phytonematodes and compromising their beneficial aspects as cover plants (Brito & Ferraz, 1987; Dias-Arieira et al., 2003; Inomoto et al., 2007). For example, U. decumbens, U. brizantha, U. humidicola, U. dyctioneura, U. ruziziensis, and the hybrid ‘grass Mulato’ (U. ruziziensis clone 44–6 × U. brizantha CIAT 6292) may increase the population of this phytonematode. It is worth mentioning that, both sorghum (Sorghum bicolor) and forage (S. bicolor × S. sudanense) are good hosts for P. brachyurus. Contrarily, black oats (Avena strigosa) are a poor host, while white oats (Avena sativa) and yellow oats (A. byzantina) are good hosts. Moreover,, few studies on the host reaction of wild turnip (R. sativus L. var. oleiferus) to P. brachyurus indicated that this crop is a bad host for the nematode. Comparably, millet (Pennisetum glaucum) and black oats are poor hosts to P. brachyurus. It means that these three plants shelter and feed P. brachyurus in their roots, allowing their reproduction, although at low levels.

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Brassicaceae residues plant residues have been reported as biofumigants after incorporation into the soil. These materials exhibit biocidal activity due to the presence of glucosinolates that can suppress soil-borne pests and diseases (Kirkegaard et al., 1993, 1998). According to the literature, Brassica spp. may drastically reduce the populations of M. incognita, M. javanica, H. schachtii, and P. neglectus (Thierfelder & Friedt, 1995; Potter et al., 1998; Monfort et al., 2007). In the Andean region of Peru, the aerial part of canola (B. napus) and mustard (Sinapis alba) plants have been used as suppressants for Meloidogyne spp. and Pratylenchus spp. in potato (Solanum tuberosum) crops, as the decomposition of the incorporated material allows the release of substances toxic to nematodes. Other examples of organic residues are sorghum (S. bicolor (L.)  (Moench, 2014)), Neem (A. indica) (Akhtar, 2000), castor bean cake (Pedroso et al., 2018) and pork beans (Canavalia ensiformis) (Rocha et al., 2017). The use of these materials has been explored in organic agriculture and they are recommended for small farmers, including family farming. The soil incorporation of industrial plant residues as well as straw to control phytonematodes in sustainable management involves quite complex and little-­ known mechanisms. The compounds produced through the biomass decomposition processes comprise an arsenal that can exhibit nematotoxic activity, induce plant defense mechanisms, as well as favor soil microfauna, increasing the fungi and bacteria populations including, among other organisms, also natural enemies. The combination of these mechanisms can hence kill or paralyze the phytonematodes. One of the examples of industrial residues that have a well-established action in the control of phytonematodes is called ‘manipueira’. This type of residue from the processing of cassava in flour mills, found in some regions of Brazil, is abundant in macro and micronutrients and, also, in cyanogenic glycosides, mainly linamarin. These compounds, when hydrolyzed, release cyanide gas, toxic to most life-forms, including nematodes. However, the amounts of hydrocyanic acid (HCN) in the ‘manipueira’ can vary with its origin, because there may be differences in cultivated cassava cultivars, storage time and type of processing, among other factors (Fonseca et al., 2018). It is worth noting that the use of cover plants can increase the organic matter content levels in soil, making it more friable and non-compacted. With the decomposition of green manures the population of nematodes is reduced, probably due to the release of different allelochemical substances. Moreover, some changes in soil physio-chemistry (pH, porosity, organic matter levels, water infiltration rate, moisture retention, macro, and micronutrients levels) may generate an effective suppression of phytonematodes in distinct types of soils and regions worldwide, when treated with organic materials (Oka, 2010). Taking into account the aforementioned aspects, it is worth to emphasize the relevance of adequate management choices, considering cover plants and the use of biopesticides based on plant residues. These eco-friendly approaches promote not only sustainable nematode control but may also interfere positively in the physio-­ chemical and microbiological soil characteristics. However, it is noteworthy that, although the management of nematodes with cover plants and plant residues is

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frequently used and effective, it has been observed that there are few recent scientific works dealing with these potentially promising materials, hindering an adequate measurement of the practical results obtained.

1.6 Animal and Agro-industrial Wastes Waste can be defined as any useless, undesirable, or disposable material resulting either after use or through a manufacturing process. These materials can be of animal origin, such as manure, feathers, bones; of vegetable origin, as agricultural waste; and of industrial origin, as by-products (Chindo et al., 2012). In this sense, Brazil stands out as being one of the largest producers in the world of agricultural and livestock wastes, available for different uses. Due to the high content of organic matter, these residues are often recycled in soil to favor agricultural productivity through fertilization. However, many materials are useful in the control of pathogens or phytonematodes. This practice has been increasingly used since it is inserted in the cyclical economic model in which waste management allows the reuse of by-products, as well as contributes to environmental well-being (Ntalli et al., 2020a). The beneficial effects of incorporating organic matter from waste in phytonematodes control include (i) increased availability of macro and micronutrients in soil; (ii) changes in the physical, chemical, and biological properties of soil; (iii) direct or indirect stimulation of the proliferation of natural enemies and (iv) increase in plant tolerance (Ntalli et al., 2020a; Peiris et al., 2020). Possible disadvantages include (i) the lack of standardization of recycled materials and/or compounds; (ii) the availability of the quantity of material needed for effective control of phytonematodes, mainly in large-scale use; (iii) accessibility to waste material and transportation to the destination of application; (iv) limitations of the application method; (v) presence of possible microbial contaminants such as fecal coliforms, bacteria (e.g., Salmonella and Escherichia coli) as well as residues containing heavy metals; (vi) application limited to the amounts allowed by local legislation; and (vii) suitability as to the timing of manure application. The use of manure is useful in the control of phytonematodes and, for this reason, has been the most studied and used in this context. The residues can be derived from livestock (bovine, equine, caprine, swine), poultry (chicken, turkey), and fish. The use of these manures can cause direct and indirect effects on phytonematodes, depending on local characteristics, such as soil and crop types, as well as local microbiota. In Brazil, the use of animal waste to control agricultural pests is an age-old practice used mainly by small producers. The country geographical and climatic conditions, favorable to both agriculture and livestock, offer ideal conditions for the practice to be increasingly applied. Currently, the growing appreciation of agricultural production, with a reduction in the use of synthetic agrochemicals has driven both research and producers to use alternative strategies that are efficient for pest

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control, especially phytonematodes, for which the use of animal waste has been one of the most common control applications. However, there are still few studies that address in-depth the properties and modes of action involved in this pest control approach. An example of a direct effect is related to the high nitrogen content and the low C:N ratio provided by manure. In general, the greater availability of N in the residue, mainly in the form of uric acid, the greater the capacity of the organic material to control nematodes (Akhtar & Malik, 2000). This is due to the formation of ammonia as one of the by-products of the organic matter decomposition. The decomposition of materials with high availability of N results in the formation of high concentrations of ammonia in the soil, which are toxic, permeating the cell membranes of phytonematodes, resulting in their inactivation or death (Peiris et al., 2020). Among the types of animal waste, poultry manure has been the most used and effective in controlling phytonematodes. According to a meta-analysis study, this type of manure provided a 64% reduction in root-knot nematodes, on average, in studies published between 2008–2018 (Peiris et al., 2020). Table 1.1 shows some examples of the action of different types of animal wastes, for phytonematodes control. Although poultry manure is more effective, in general, some studies also showed beneficial effects in the use of livestock manure (Table 1.1). Such action is generally due to indirect effects, such as the changes in physical, chemical, and biological properties of soil (pH, microflora, and microbiota, for example), as well as the induction of resistance in certain types of crops. Hence, in most cases, the use of animal manure increases productivity and product quality. Although the mode of action is still not very clear, there is strong evidence that the increase in the number of free-living nematodes together with the increase in the population of fungi, bacteria, and nematode predatory mites are responsible for the reduction of phytonematodes. El-Marzoky et  al. (2018) evaluated the use of cow, horse, and turkey manure (30–40 kg/tree), in a Balady mandarin orchard contaminated with T. semipenetrans, Pratylenchus spp., Tylenchorhynchus spp., and Helicotylenchus spp. A greater density reduction for the four phytonematodes was observed when turkey (75–78%), horse (64–68%), and cow manure (53–57%) were used, respectively. One of the main factors responsible for this effectiveness appears related to the nitrogen content of the applied residues. According to the authors, the nitrogen content in turkey, horse, and cow dung is about 29,100 ppm, 19,500 ppm, and 17,700 ppm, respectively. For this reason, greater effectiveness in the use of turkey manure was noted, even when it was applied in smaller amounts than in the other treatments. Other types of waste that have been evaluated for use against phytonematodes are agro-industrial by-products. Among them, some commonly used ones are fruit and cereal peels (Ebrahimi et al., 2016; Maleita et al., 2017; Ali & Zewain, 2018; Izuogu et al., 2019), date palm fibers (Montasser et al., 2016), maize silage (Westphal et  al., 2016), brewers spent grain (Thligene et  al., 2019); dry-grape marc (Nico et al., 2004), fish waters (El-Deeb et al., 2019), garlic straw (Gong et al., 2013),

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Table 1.1  Examples of studies involving animal waste for the control of phytonematodes Rate of Waste types application Phytopathogen Cattle dung 100 mL P. zeae (compost tea) (100%)

Experimental resultsb Strategya FE (on maize) NPR (77.1%)

Cattle slurry

48.2 t/ha

G. rostochiensis

PE (on potato) EMR (85.0%)

Cattle slurry

48.2 t/ha

G. pallida

PE (on potato) EMR (82.0%)

Chicken manure Chicken manure Chicken manure Chicken manure Chicken manure Chicken manure Chicken manure Chicken manure Chicken manure

30 kg/tree

FE (in fruit orchard) M. incognita GE (on cucurbit) H. schachtii FE (on sugarbeet) Pratylenchus spp. FE (in fruit orchard) Tylenchorhynchus FE (in fruit spp. orchard) Hoplolaimus spp. FE (in fruit orchard) Helicotylenchus FE (in fruit spp. orchard) M. incognita PE (on melon)

3 g/plant 40 t/ha 30 kg/tree 30 kg/tree 30 kg/tree 30 kg/tree 3 t/ha

T. semipenetrans

2.5 kg/m2

Meloidogyne incognita

Cow dung

20 t/ha

M. incognita

Cow dung

20 g/kg soil

M. incognita

Cow dung

60 t/ha

H. schachtii

Cow manure

100 kg/ha

M. incognita

Goat dung

20 t/ha

M. incognita

Goat manure

3 g/plant

M. incognita

Goat manure

800 kg/ha

M. incognita

FE (on eggplant cv. Baladi) FE (on sweet potato)

NPR (68.5%) NGR (61.0%) NPR (92.4%) NPR (71.4%) NPR (80.1%) NPR (83.0%) NPR (61.9%) NPR (87.3%) NPR (81.2%); NGR (63.0%); EMR (70.0%) NPR (76.8%)

References Izuogu and Usman (2019) George et al. (2016) George et al. (2016) El-Metwally et al. (2019) El-Deeb et al. (2018) Nasresfahani (2017) El-Metwally et al. (2019) El-Metwally et al. (2019) El-Metwally et al. (2019) El-Metwally et al. (2019) Abdel-Dayem et al. (2012) Osman et al. (2018)

Osunlola and Fawole (2015) PE (on RGR (92.2%); Zafair et al. Ammimajus) EMR (81.5%); (2018) NPR (91.5%) FE (on sugar NPR (67.9%) Nasresfahani (2017) beet plants cv. 005) PE (on okra) NPR (76.6%) Tanimola and Akarekor (2014) FE (on sweet NPR (82.1%) Osunlola and potato) Fawole (2015) GE (on RGR (56.6%) El-Deeb et al. cucurbit) (2018) GE (on RGR (89.0%) Pakeerathan tomato) et al. (2009) (continued)

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Table 1.1 (continued) Experimental resultsb NPR (67.8%)

Waste types Goat manure

Rate of application Phytopathogen 100 kg/ha M. incognita

Horse dung

20 t/ha

M. incognita

FE (on sweet potato)

NPR (74.6%)

Horse manure (composted) Pig slurry

10 g/kg

M. incognita

GE (on tomato)

RGR (4.7%)

24.7 t/ha

G. Rostochiensis

PE (on potato) EMR (77.5%)

Pig slurry

24.7 t/ha

G. pallida

PE (on potato) EMR (75.0%)

Pig manure

100 kg/ha

M. incognita

PE (on okra)

P. zeae

FE (on maize) NPR (86.4%)

M. incognita

FE (on sweet potato)

NPR (87.3)

M. incognita

PE (on okra)

NPR (86.7%)

100 mL Poultry (100%) dropping (compost tea) Poultry dung 20 t/ha

Poultry manure

100 kg/ha

Strategya PE (on okra)

NPR (60.0%)

References Tanimola and Akarekor (2014) Osunlola and Fawole (2015) Siddiqui and Akhtar (2008) George et al. (2016) George et al. (2016) Tanimola and Akarekor (2014) Izuogu and Usman (2019) Osunlola and Fawole (2015) Tanimola and Akarekor (2014)

FE field experiment, GE greenhouse experiment, PE potted experiment NPR Nematode Population Reduction, RGR Root Galls Reduction, EMR Egg Masses Reduction

a

b

onion bulb (Youssef & El-Nagdi, 2010), sawdust (Siddiqui & Akhtar, 2008; Hassan et al., 2010; Faruk, 2019), oil-seed cakes (Pedroso et al., 2018; Faruk, 2019; Pandeya et al., 2019), olive mill waste (Cayuela et al., 2008), rice bran (Hassan et al., 2010; Faruk, 2019) and wine industry by-products (Reiner et al., 2016). Brazil has several territorials, climatic, and productive advantages that favor the possibility of using agricultural and industrial wastes for pest control. The following passage describes a successful case in the development of control measures against phytonematodes. The liquid residue (100 mL) resulting from the extraction of sisal fibers (Agave sisalana) was evaluated against Radopholus similis on Grand Naine banana trees under greenhouse conditions. Compared with water-treated plants, the fresh (FR1) and fermented (FR2) residues in concentrations of 25% caused a reduction in the number of juveniles in soil (FR1 = 66% and FR2 = 80%) and in roots (FR1 = 84 and FR2 = 77%). The nematicidal effect of such residues may be associated with secondary metabolites, such as alkaloids, saponins, terpenes, tannins, flavonoids and glycosides. There are reports that the saponins in this type of residue

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may interact with nematode cuticle proteins, leading to their disruption (Jesus et al., 2015). Sugar beets filter cake or mud is an industrial waste evaluated under field conditions in bananas (Cv. Williams) infected with M. incognita. The application of 1 ton/ feddan provided a gall reduction of 74.1%, 56.8%, and 79.6% in the vegetative, flowering, and harvesting stages, respectively. The observed action likely resulted from direct and indirect factors such as the release of nematotoxic compounds from the decomposition of organic matter as well as the activity of certain antagonistic or predatory nematodes. In addition to the nematotoxic effect, the residues provided an increase of 28.4% (in tons) in banana yield per feddan (Youssef & El-Nagdi, 2010). A dry cork compound, obtained as industrial residue and submitted to the standard composting process, showed a nematotoxic activity evaluated in tomato plants parasitized by M. incognita race 1 and M. javanica. In a growth chamber, the residue provided, at concentrations of 25–100% (v/v), a reduction in the roots gall index of 27.3–54.5% and 27.5–75.0%, when compared to the untreated control. A similar and considerable reduction in the final nematode population of M. incognita (65.3–98.5%) and M. javanica (65.5–99.7%) was also observed. As the experiment was conducted on sterile soil, the nematotoxic effects observed are probably related to the compounds present in the composted material, such as tannins and/or phenolic compounds. These types of cork constituents have already shown suppression of nematodes as organic amendments (Nico et al., 2004). Aqueous extracts of agro-industry coffee-silverskin waste were evaluated in tomato plants infected with M. incognita. In a greenhouse, each potted plant received 1 L of these extracts, at a 50% concentration. The residue extract reduced the number of eggs and juveniles/g of root by 50.9% and 44.4% respectively, compared to the control. The final nematode population density was reduced by 47.8%. The nematotoxic action observed was attributed to the release of polyphenols (caffeoyl and feruloyl quinic derivates) from the coffee epidermis (Thligene et al., 2019). Despite the many successful examples in the use of residues to control phytonematodes, few products are available on the market. In the case of animal waste, these are often purchased from breeding sites close to the cultivation area or the breeder himself supplies the animal waste to the crop production area. As for agro-­ industrial waste, these can be purchased directly from the productive sector, since industries generally dispose of or make the material available at affordable prices. It is worth to mention that, in both situations, greater accessibility to waste is a favorable factor for acquisition and, therefore, its use on a smaller scale. In addition, for both producers and industries, disposing of the respective residues for use in agriculture, for example, is a viable way to treat residues and an opportunity to reduce production costs. Thus, there is a strong trend in expanding the use of animal and agro-industrial wastes in pest control. Their use in the control of phytonematodes has multiple benefits, such as the reduction in the application of synthetic nematicides combined with an increase in crop yields. Consequently, the reduction of environmental impact is two-fold, through recycling the material that would be discarded, and/or by reducing the contamination that may result from the use of conventional agrochemicals. On the other hand, there is a need to ensure reliability

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in the use of waste for food production. In this sense, the use of methods of detecting pathogenic organisms for human and animal health in said residues, for example, may be appropriate. It is worth mentioning that cost-benefit is also a factor to be considered.

1.7 Blends A current trend is to mix compounds to get around a specific situation, for example, pest control. Such a mixture can be called a blend, in which two or more substances, when mixed together, result in a synergic, superior effect than when each one is applied alone. In the case of phytonematodes control, several strategies of mixtures and combinations of natural compounds have been studied. Figure  1.2 presents some examples of mixtures evaluated against several types of phytonematodes, showing that animal manure and/or microorganisms, such as fungi and bacteria, are included in most combinations. In general, a single biocontrol agent is used against a specific pathogen. On the other hand, mixing or combining different sources of bioactive ingredients can guarantee the multifunctionality of the resulting product. In this way, multiple disease suppression mechanisms may be implemented simultaneously for each biocontrol approach applied (Siddiqui & Akhtar, 2008; Udo et al., 2020). Depending on the mode of action of each constituent, the effect of their combination may result in synergism or cause an additive effect when compared to individual applications (Asif et al., 2017b; Cookey et al., 2019). In this way, the mixture of natural products can generate marked effects in terms of pest control as well as favoring productivity. Consequently, greater effectiveness can be achieved when applying suitable combinations. However, the use of combinations requires care, since antagonistic effects can occur. In the case of concomitant use of microorganisms, for example, the competitiveness and possible interferences that one species may cause to another one must be evaluated. In addition, it is of fundamental importance to evaluate the ideal dosage of each substance and the method of application to avoid phytotoxicity. In fact, combinations also require a correct timing, since non-simultaneous application can potentiate the positive effects. As shown in Fig. 1.2, there are several possibilities for mixing different compounds. In the case of mixing compounds, it becomes even more complex and challenging to identify the mechanism of action involved. For this reason, despite their increasing uses, few studies have addressed in-depth the main ingredients active in the control of phytonematodes, present in mixtures. Moreover, few studies have sought to understand the mode of action involved when more than one asset is present. In general, hypotheses are derived from the individual effects already established. Most of the time, the studies only seek to verify effectiveness in terms of pest control and increased productivity. The following citations show some successful

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Fig. 1.2  Venn diagram indicating some examples of mixtures evaluated against several types of phytonematodes. The numbers indicated in the intersections correspond to the citations: 1 (Abdel-­ Monaim et al., 2018); 2 (D’Addabbo et al., 2011); 3 (Asif et al., 2017b); 4 (Asif et al., 2017a); 5 (Buena et al., 2007); 6 (Cole et al., 2020); 7 (Cookey et al., 2019); 8 (Djiwanti et al., 2019); 9 (El-Deeb et al., 2019); 10 (El-Nagdi et al., 2019); 11 (Gong et al., 2013); 12 (Gupta et al., 2019); 13 (Mehtab et al., 2013); 14 (Meyer et al., 2011); 15 (Meyer et al., 2015); 16 (Mostafanezhad et al., 2014); 17 (Nath & Singh, 2011); 18 (Rao et al., 2017); 19 (Ravindra et al., 2014); 20 (Renčo & Kováčik, 2015); 21 (Rizvi et al., 2015); 22 (Rostami et al., 2014); 23 (Sari et al., 2018); 24 (Siddiqui & Akhtar, 2008); 25 (Tiyagi & Ajaz, 2004); 26 (Udo et al., 2020); 27 (Varkey et al., 2018). (Author: Lívia Cristina de Souza Viol)

cases using the combination of natural products for the phytonematodes control, together with the supposed mode of action involved. In an experimental field, the nematotoxic effect of oilseed cakes of neem (A. indica), castor (R. communis), groundnut (A. hypogaea), linseed (L. usitatissimum), and sunflower (H. annuus) was achieved when chickpea seeds (C. arietina) cv. K-850 were pre-treated with the fungus Paecilomyces lilacinus. The application of 110 kg/ha of each oilseed cake in soil contaminated with several species of phytonematodes (Hoplolaimus indicus, Helicotylenchus indicus; Rotylenchulus reniformis; T. brassicae; Tylenchus filiformis; M. incognita; Hemicriconemoides sp.; Longidorus elongatus, Xiphinema basiri and Trichodorus mirzai) showed a reduction in their populations between 70.1% and 75.1% and between 78.7% and 83.6%,

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in absence and presence of P. lilacinus, respectively, when compared to untreated plants. All plants submitted to the dual treatment showed improvement in all growth parameters. Among them, Neem cake + P. lilacinus increased by 389.0% total plant weight and by 279.7% the number of pods per plant, compared to untreated controls. These beneficial effects are attributed to the release of nematotoxic substances and nutrients, such as sources of N, by the organic amendments. In the case of P. lilacinus, this fungus is capable of colonizing and destroying female reproductive organs, eggs, and nematode cysts. Its multiplication may have been favored by the cake substrate added to the soil, which further enhanced its effect, increasing plants resistance to the nematodes attack (Tiyagi & Ajaz, 2004). In another study, organic wastes (biosolids, horse manure, neem leaf litter, and sawdust) were tested in the presence and absence of the fungi Glomus intraradices (G) combined or not with the rhizobacterium Pseudomonas putida (P). The greenhouse experiments with tomato plants infected with M. incognita in pots containing 1 kg of soil were carried out using 10 g of organic waste and 10 mL of the suspension (1.5 × 107 cells/mL in nutrient broth) of each microorganism. The combination of the three constituents (organic waste + G + P) favored higher efficiencies regardless of the type of waste used. Among them, neem leaf litter was the most effective in the presence of the two microorganisms and provided a reduction of 46.5% and 17.4% in the number of galls and nematodes, respectively. Such combination also contributed to an increase in length (110%) and shoot dry weight (137%) of tomato plants, compared to controls. In the case of G, its inhibitory action may be associated with the activation of plant defense mechanisms and/or with an increase of the content in phosphorus, phenylalanine, and serine in plant roots. As for P, in addition to improving plant growth, it can suppress pathogens by the synthesis of enzymes, capable of modulating plant hormone levels, and antibiotics, capable of killing pathogens. Concerning organic residues, they are known to improve the soil structure by supplying nutrients and accumulating antagonistic organisms. Among them, the best efficiency of neem leaf litter appeared related to their highest C:N ratio (19:8) (Siddiqui & Akhtar, 2008). Experiments with tomato plants parasitized by M. incognita were carried out in containers including 11  kg of incorporated soil. Among some types of substances evaluated, the combination involving 1.5 kg of rabbit manure and 220 g of raw garlic straw and coverage with polyethylene film was the most efficient and capable of reducing the galling index by 72.3%, increasing tomato production by 72.6%. GC-MS analysis performed with garlic straw indicated that volatile sulfur-­containing compounds, such as dimethyl disulfide, dipropyl disulfide, and diallyl disulfide, were mainly responsible for the inhibition of M. incognita by this residue. In addition, it was observed that the use of raw garlic straw disturbed the reproduction of M. incognita as it caused a sharp reduction of female numbers, in addition to causing a delay in the life cycle of the nematode. The incorporation of rabbit manure and plastic film into the garlic straw dramatically contributed to the bio-fumigant efficacy. Rabbit manure may have facilitated the decomposition of garlic straw and, consequently, contributed to the increase in the emission of bioactive compounds, in addition to releasing nematotoxic by-products. In the case of the plastic film, it may have prevented the escape of nematicidal volatiles, also serving as a physical barrier (Gong et al., 2013).

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Vermicomposts of different types of animal wastes (cow, buffalo, sheep, horse, and goat manure) prepared with neem oil, the aqueous leaf extract of custard apple (Annona squamosa), or aqueous extract of the garlic bulb, singly and binary combined with agro wastes (gram bran, straw, wheat bran, barley bran, and rice bran) were evaluated on tomato in a field infected with a lesion nematodes, Pratylenchus sp. Several of the evaluated combinations significantly reduced the nematode population, with emphasis on the mixtures of vermicompost with garlic extract, neem oil, and custard apple extract, which reduced the nematodes by at least 93.8%, 92.0%, and 84.6%, respectively. In addition to nematode control, the use of combinations allowed early flowering and improved both growth and productivity, the last item being quadrupled, compared to control. A large amount of humic acid, resulting from the vermicomposting process, together with the diversity of macro and micronutrients, vitamins, enzymes, and growth hormones contributed to the increase in the tomato productivity, while the constituents of the plant extracts used in the combinations affected the phytonematodes nervous system (Nath & Singh, 2011). Due to the potentiation of the effect when the use of constituents is combined, the tendency to develop new mixtures is increasing. There are several products on the market involving mixtures available for use in pest control. So far, most products involving mixtures for use against phytonematodes showed the presence of microorganisms. This fact can be a result of both the effectiveness and the existence of a legislation favorable to its commercial approval. In this sense, it is of fundamental importance that regulations are developed for the approval of other natural resources, such as extracts, also useful for pest control. In addition, the use of mixtures is a very attractive factor for both small, medium, and large producers, since it facilitates their joint application. Despite its great potential in pest control, there is a need of further data about mixtures. One of the challenges for the development of blends or combinations of natural compounds is to identify the active principles involved when the constituents are applied together. In addition, data are lacking on the ideal dosage, the control of possible contaminating agents (microorganisms, pesticides and heavy metals), adequate application time, duration, and mode of action. It is worth mentioning that, in the case of natural products, the variability of the materials can interfere with the final result. To minimize such limitations, some strategies can be adopted such as dosage standardization, forms of application, measures to evaluation pest control effects, among others.

1.8 Challenges and Future Trends Market globalization offers the consumer a great diversity of products that can be deferred by several factors such as quality, price, safety, impacts on the environment, among others. To ensure consumer reliability, norms and standards have been established by government agencies. They aim at formalizing and institutionalizing the minimum requirements of standard criteria, to obtainin products with greater

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competitiveness, safety, and general quality. In recent decades, together with legal requirements, the demand for safer and high-quality food has been growing. These actions have been reflected in the implementation of increasingly strict safety regulations, so that standards related to product quality have been continuously raised higher. Sustainable agriculture needs pest control methods that have a low impact on the environment, as well as human and animal health. Thus, there is a tendency to use resources of natural origin since several formulated products, for example, synthetic nematicides, have been withdrawn from the market due to toxicity. However, there is still a lack of more sustainable and effective commercial products to control the many pests. Figure 1.3 shows the main trends and challenges for the development of new, more sustainable pesticides.

Fig. 1.3  Main trends (left) and challenges (right) for the development of new, more sustainable pesticides. (By Lívia Cristina de Souza Viol)

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A successful research and development process that leads to a commercial product must meet several criteria, especially biological, environmental, toxicological, regulatory, and commercial. Therefore, they need to be planned in a careful and standardized way, starting from the origin of the product. In this case, the choice of raw materials that enable their sustainable use is a great challenge, as their availability and accessibility must also be taken into consideration. Products based on natural sources such as plant extracts may involve renewable resources. However, the growing world demand for this type of product requires large-scale production and an increase in cultivated areas. In addition, seasonality, according to water and nutritional needs, the photoperiod, and other particularities of each botanical species may limit its production. Thus, it is necessary to use different sources, to promote the diversification of the natural products used, according to the specific characteristics of each region. Another challenge is the quality control. In this case, the lack of adequate standards and analytical procedures for the manufacture of homogeneous batches, free from contamination (chemical or microbial) and which ensure product shelf-life are still factors to be overcome. In addition, it is important to emphasize the need to optimize methods that use eco-friendly technology, since most conventional extraction processes use solvents, and toxic processes that cause disposal problems, with potential environment contamination. Another relevant effort is the supply of products of adequate quality and quantity, at low cost to farmers, worldwide. One of the ways to overcome these limitations is the involvement of a multidisciplinary team (composed of a nematologist, agronomist, biochemist, chemist, statistician, plant physiologist, environmental toxicologist, biologist, among others) throughout the production process. Thus, it becomes possible to accelerate and standardize the procurement of raw material in a controlled manner to establish formulations that maintain the stability of products, using technologies that have little or no impact on human and animal health, as well as on the environment. Another way to streamline the development, manufacture, and commercialization of environment friendly alternatives is through the cooperation between the public and private sectors. In this context, the discovery of new substances and research on formulations linked to their production and distribution would boost the natural products commercialization and use. In addition, investing heavily in product quality and certification is also an important market strategy. An important point for product development is the characterization of its toxicity. In this case, the natural resources selected for pest control must be specifically active against the targeted pest. Therefore, the risk of affecting non-targeted organisms (animals, beneficial insects, and humans) should be relatively low. For this, it is necessary to carry out standardized safety tests, as required in regulations for product registration, so that permission to launch products based on natural resources is granted. In addition to toxicological assessments, the guidelines established by several countries, such as the USA, Canada, the European Union and others for product registration include important information such as (i) identification, physical-­ chemical characterization and analysis of chemical compounds/complexes and/or

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formulated product; (ii) manufacturing methods using standardization criteria that ensure the reproducibility of the physical-chemical characteristics and biological properties of the product; (iii) waste analysis; and (iv) suitability of packaging, labels, and package inserts. In general, quality regulatory mechanisms aim at increasingly meet the demand of society, the media, and national/international trade, which currently calls for products that have a low impact on the environment and result in safe food. However, in several countries (mainly in developing regions), a greater incentive to utilize these products is necessary, with the establishment of guidelines and less bureaucratic processes. Moreover, it is also necessary to encourage and inform the farmers about the use of natural products in the management of agricultural pests. Another growing trend is the use of products of natural origin together with integrated management and in organic agriculture. The tendency for the coming years concerning the research programs on sustainable management of plant-parasitic nematodes, invariably should use to some extentand in association or not, the alternatives pointed out below. Currently, there is an increase in the number of research groups and companies that are investing in pest control, including phytonematodes, following a rationale of “green” technologies. In this context, research groups and companies that invest in quality control systems will stand out, by obtaining natural products or by-­ products that are more homogeneous and free of contaminants. Besides the standardization of the conditions for obtaining the aforementioned materials, differentiating factors are preparing and formulating, as well as, characterizing products to guarantee duplication, reproduction, reliability, stability, efficiency, safety, and customer satisfaction. The use of nanomaterials generated from green chemistry strategies represents a solution increasingly used in the industry, medicine, food, and agriculture sectors. In this case, nanocarriers such as metallic or polymeric nanoparticles and nanoemulsions can carry compounds, extracts, or blends resulting in a more controlled release and in a lesser amount of material applied. Blends obtained from combinations such as (i) different plant extracts, (ii) plant extracts containing microorganisms, (iii) plant extracts associated with biofertilizers, among others, are another tendency for phytonematodes control. Moreover, this strategy can be used multifunctionally for the control of different pests as well as for plant development. Regarding IPM, there is a tendency for this alternative to absorb strategies such as nanotechnology and blends, following “green” chemistry recommendations, making pest control more efficient and reducing its impact. In addition, a new trend is the holistic use of IPM that seeks to further integrate the farmer into the process of designing solutions for pest control and minimizing socioeconomic and environmental impacts. To that end, stakeholders should, in support of researchers and policymakers, be more aware of the need to adopt botanical pesticides and other natural products as safe tools for pest control. Researchers and scientists working with these products are tasked with providing consistent and reproducible field effectiveness data.

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Consequently, the increased use of eco-friendly products in pest control, together with IPM strategies will lead to better acceptability of agricultural products in niche markets and increased customer satisfaction. Moreover, this may also contribute to improve international trade, as well as biodiversity conservation, environmental protection, food security, and human health.

1.9 Conclusion Several research successes, regarding the use of natural and residual resources, show efficient control of different species of phytonematodes in distinct crops. In many cases, various extracts, essential oils, and residues, as well as their mixture as blends, can also increase productivity and contribute, to some extent, to human, animal and environmental health, ultimately boosting the bioeconomy. Furthermore, cover crop plants can also be used in association with previous strategies. Although there are still only a few natural-based nematicidal products available and regulated on the market, efforts to adapt and make them available have been growing, both by public and private institutions and government agencies. It is worth mentioning that many limitations in the availability of these resources, mainly on a larger scale, still exist, such as their accessibility, standardization, and adequate dosages. In addition, there is also a need for more in-depth studies to assess the effectiveness of these resources in the short and long term. In this sense, the realization of field experiments, identification of active molecules together with the development of suitable formulations for application, will certainly contribute to increase the availability of these resources for all types of producers. Finally, there is a tendency to integrate strategies such as nanotechnology, the use of blends, the IPM as well as Holistic Pest Management that together can guarantee food sovereignty, well-paid work for agricultural workers, and food security for the growing world population.

References Abbassy, M.  A., Abdel-Rasoul, M.  A. M., Nassar, A.  M. K., & Soliman, B.  S. M. (2017). Nematicidal activity of silver nanoparticles of botanical products against root-knot nematode, Meloidogyne incognita. Archives of Phytopathology and Plant Protection, 50, 909–926. Abdel-Monaim, M., Abdel-Bast, S., & Wahdan, R. (2018). Effectiveness of some organic fertilizers and bio-control agents for controlling root-knot nematodes Meloidogyne spp. in cowpea forage (Vigna unguiculata) in New Valley, Egypt. Egyptian Journal of Phytopathology, 46, 143–164. Abdel-Dayem, E. A., Erriquens, F., Verrastro, V., Sasabekki, N., Mondelli, D., & Cocozza C. (2012). Nematicidal and fertilizing effects of chicken manure, fresh and composted olive mill wastes on organic melon, Institute of Parasitology. Helminthologia, 49(4), 259–269. https://doi.org/10.2478/s11687-012-0048-4

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Akhtar, M. (2000). Nematicidal potential of the neem tree Azadirachta indica (A. Juss). Integrated Pest Management Reviews, 5, 57–66. Akhtar, M., & Malik, A. (2000). Roles of organic soil amendments and soil organisms in the biological control of plant-parasitic nematodes: A review. Bioresource Technology, 74, 35–47. Akyazi, F. (2014). Effect of some plant methanol extracts on egg hatching and juvenile mortality of root-knot nematode Meloidogyne incognita. American Journal of Experimental Agriculture, 4, 1471–1479. Ali, S.  F., & Zewain, Q.  K. (2018). Efficiency of some bio and organic agents to control root knot nematode Meloidogyne spp on cucumber plants Cucumis Sativus. Tikrit Journal For Agricultural Sciences, 18, 117–127. Andrés, M. F., González-Coloma, A., Sanz, J., Burillo, J., & Sainz, P. (2012). Nematicidal activity of essential oils: A review. Phytochemistry Reviews, 11, 371–390. Aoudia, H., Ntalli, N., Aissani, N., Zaidi-Yahiaoui, R., & Caboni, P. (2012). Nematotoxic phenolic compounds from Melia azedarach against Meloidogyne incognita. Journal of Agricultural and Food Chemistry, 60, 11675–11680. Asif, M., Khan, A., Tariq, M., & Siddiqui, M. A. (2017a). Sustainable management of root knot nematode Meloidogyne incognita through organic amendment on Solanum lycopersicum L. Asian Journal of Biology, 1, 1–8. Asif, M., Tariq, F. A. M., Khan, A., Ansari, T., Khan, F., & Siddiqui, A. M. (2017b). Potential of chitosan alone and in combination with agricultural wastes against the root-knot nematode, Meloidogyne incognita infesting eggplant. Journal of Plant Protection Research, 57, 288–295. Bakkali, F., Averbeck, S., Averbeck, D., & Idaomar, M. (2008). Biological effects of essential oils a review. Food and Chemical Toxicology, 46, 446–475. Barbosa, A., Silva, L., Ferraz, C. M., Tobias, F. L., et al. (2019). Nematicidal activity of silver nanoparticles from the fungus Duddingtonia flagrans. International Journal of Nanomedicine, 14, 2341–2348. Bills, G.  F., & Gloer, J.  B. (2016). Biologically active secondary metabolites from the fungi. Microbiology Spectrum, 4, 10. Retrieved from https://doi.org/10.1128/microbiolspec. FUNK-­0009-­2016 Bogner, C. W., Kamdem, R. S., Sichtermann, G., Matthäus, C., et al. (2017). Bioactive secondary metabolites with multiple activities from a fungal endophyte. Microbial Biotechnology, 10, 175–188. Bömke, C., & Tudzynski, B. (2009). Diversity, regulation, and evolution of the gibberellin biosynthetic pathway in fungi compared to plants and bacteria. Phytochemistry, 70, 1876–1893. Brito, J.  A., & Ferraz, S. (1987). Seleção de gramíneas antagonistas a Meloidogyne javanica. Nematologia Brasileira, 10, 260–269. Buena, A. P., García-Álvarez, A., Díez-Rojo, M. A., Ros, C., Fernández, P., Lacasa, A., & Bello, A. (2007). Use of pepper crop residues for the control of root-knot nematodes. Bioresource Technology, 98, 2846–2851. Butnariu, M., & Sarac, I. (2018). Essential oils from plants. Journal of Biotechnology and Biomedical Science, 1, 35–43. Caboni, P., & Ntalli, N. (2014). Botanical nematicides, recent findings. In A. D. Gross, J. R. Coats, S.  O. Duke, & J.  N. Seiber (Eds.), Biopesticides: State of the art and future opportunities (pp.  145–157). ACS Division of Agrochemicals. Retrieved from https://doi.org/10.1021/ bk-­2014-­1172 Calvo, A. M., Wilson, R. A., Bok, J. W., & Keller, N. P. (2002). Relationship between secondary metabolism and fungal development. Microbiology and Molecular Biology Reviews, 66, 447–459. Cayuela, M. L., Millner, P. D., Meyer, S. L. F., & Roig, A. (2008). Potential of olive mill waste and compost as biobased pesticides against weeds, fungi, and nematodes. Science of the Total Environment, 399, 11–18. Chindo, P. S., Bello, L. Y., & Kumar, N. (2012). Utilization of organic wastes for the management of phyto-parasitic nematodes in developing economies. In S. Kumar & A. Bharti (Eds.),

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Tiku, A. R. (2020). Antimicrobial compounds (Phytoanticipins and Phytoalexins) and their role in plant Defens (pp. 845–868). Springer. Tiyagi, S. A., & Ajaz, S. (2004). Biological control of plant parasitic nematodes associated with chickpea using oil cakes and Paecilomyces lilacinus. Indian Journal of Nematology, 34, 44–48. Udo, I.  A., Uko, A.  E., & Etim, D.  O. (2020). Management of root-knot disease in okra with poultry manure and leaf extracts of Senna alata. South Asian Journal of Parasitology, 4, 1–10. Umar, I., Muhammad, Z., & Okusanya, B. A. O. (2010). Effect of organic amendments on the control of Meloidogyne javanica (Kofoid and White, 1919) Chitwood, 1949, on tomato (Lycopersicon lycopersicum, Mill). Agriculture, Business and Technology Journal, 8, 63–77. United States Environmental Protection Agency (USEPA). (2009). Registering pesticides, 2009. Retrieved from http://www.epa.gov/pesticides/regulating/re-­gistering/index.htm VanderMolen, K. M., Raja, H. A., El-Elimat, T., & Oberlies, N. H. (2013). Evaluation of culture media for the production of secondary metabolites in a natural products screening program. AMB Express, 3, 71. Varkey, S., Anith, K. N., Narayana, R., & Aswini, S. (2018). A consortium of rhizobacteria and fungal endophyte suppress the root-knot nematode parasite in tomato. Rhizosphere, 5, 38–42. Vinale, F., & Sivasithamparam, K. (2020). Beneficial effects of Trichoderma secondary metabolites on crops. Phytotherapy Research: PTR, 34, 2835–2842. Wang, K. H., Sipes, B., & Schmitt, D. (2002). Crotalaria as a cover crop for nematode management: A review. Nematropica, 32, 35–57. Wang, Q., Li, Y., Klassen, W., & Handoo, Z. (2007). Influence of cover crops and soil amendments on okra (Abelmoschus esculentus L.) production and soil nematodes. Renewable Agriculture and Food Systems, 22, 41–53. Westphal, A., Kücke, M., & Heuer, H. (2016). Soil amendment with digestate from bio-energy fermenters for mitigating damage to Beta vulgaris sub spp. by Heterodera schachtii. Applied Soil Ecology, 99, 129–136. Yang, Z. S., Li, G. H., Zhao, P. J., Zheng, X., et al. (2010). Nematicidal activity of Trichoderma spp. and isolation of an active compound. World Journal of Microbiology and Biotechnology, 26, 2297–2302. Yang, Z., Yu, Z., Lei, L., Xia, Z., et  al. (2012). Nematicidal effect of volatiles produced by Trichoderma sp. Journal of Asia-Pacific Entomology, 15, 647–650. Youssef, M. M. A., & El-Nagdi, W. M. A. (2010). Effect of certain organic materials in controlling Meloidogyne incognita root-knot nematode infesting banana. Archives of Phytopathology and Plant Protection, 43, 660–665. Zafair, A., Sumbul, A., & Mahmood, I. (2018). Control of root knot disease in Ammi majus using organic amendments. The Journal of Indian Botanical Society, 97, 99–107.

Chapter 2

Organic Nematicides: A Green Technique and Its Overview for Nematode Pest Management Faryad Khan, Mohammad Shariq, Mohd Asif, Taruba Ansari, Saba Fatima, Arshad Khan, Mohd Ikram, and Mansoor Ahmad Siddiqui

Abstract  Plant-parasitic nematodes (PPN) are considered a global problem and major obstacle in agriculture. They alter the histology and physiology of plants, affecting the metabolic activity of vegetative and reproductive parts thereby resulting in significant agricultural yield losses. The application of chemical nematicides is a key factor in controlling PPNs and is reported to cause serious threats to the ecosystem. Presently, the increasing interest of farmers in protecting the environment and human health has prompted research on sustainable approaches that require fewer synthetic nematicides, and fertilizers. Several methods, including organic materials, are an effective approach for controlling diseases due to PPN in field crops. Hence, this chapter focuses on the comprehensive and practical use of some plants and their by-products, organic manures, agro-industrial wastes, de-­ oiled cakes, biochar, and chitosan in reducing nematode population levels in global cropping systems. The mode of action of organic nematicides is described in terms of controlling PPNs and stimulating plant growth. These practices can be employed to manage nematodes successfully in limited resource farming systems, to achieve sustainable and eco-friendly crop productions. Keywords  Biochar · Chitosan · Chemical nematicides · Eco-friendly organic nematicides · Plant-parasitic nematodes

F. Khan (*) · M. Shariq · T. Ansari · S. Fatima · A. Khan · M. Ikram · M. A. Siddiqui Section of Plant Pathology and Nematology, Department of Botany, Aligarh Muslim University, Aligarh, India M. Asif Department of Pharmacognosy, Pharmacopoeia Commission for Indian Medicine and Homeopathy, Ghaziabad, Uttar Pradesh, India © The Author(s), under exclusive license to Springer Nature Switzerland AG 2022 K. K. Chaudhary, M. K. Meghvansi (eds.), Sustainable Management of Nematodes in Agriculture, Vol.1: Organic Management, Sustainability in Plant and Crop Protection 18, https://doi.org/10.1007/978-3-031-09943-4_2

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F. Khan et al.

2.1 Introduction Nematodes are multicellular, microscopic, and bilaterally symmetrical animals that belong to Ecdysozoa. They may vary from 250 μm to 12 mm in length and 15 to 35 μm in width. In this Phylum, plant-parasitic nematodes (PPN) encompass about 10% of the identified species, with more than 4100 taxa (Decraemer & Hunt, 2006). Many PPN are serious agricultural pests because of their short life-cycle, high population densities, and reproductive potential (Sikora & Fernandez, 2005). They damage almost every plant on the planet, including cereals, vegetables, fruits, foliage, ornamentals, and forest crops, targeting any part of the plant, including roots, stems, leaves, and flowers. Among all PPN, the most deleterious genera are Meloidogyne, Globodera, Heterodera, Pratylenchus, Helicotylenchus, Tylenchulus, Ditylenchus, Radopholus, Xiphinema, Bursaphelenchus and Rotylenchulus (Jones et al., 2013). In India, over 600 plant-parasitic species are associated with more than 700 host plants. Globally, many attempts have been made to determine the crop losses caused by PPN. Nicol et al. (2011) estimated the crop losses to world agriculture in terms of the economic value of about 80–118 billion US dollars per year. In India, agriculture faces a decline in yield encompassing 19.6%, reaching up to 60% in most severe cases (Gowda et al., 2017). The annual average losses due to PPNs in India was estimated to be Rs. 210 crores (Jain et al., 2007). Meloidogyne incognita is one of the most severe soil PPN, causing financial losses accounting for around $100 billion per year, worldwide (Trudgill & Blok, 2001). Other economically significant PPNs are G. rostochiensis on potato, Radopholus similis on banana, T. semipenetrans on citrus, Bursaphelenchus cocophilus on coconut, and Rotylenchulus reniformis on legumes. Generally, these economically important nematodes are managed by applying synthetically-derived nematicides. However, the rising riss and impact caused to the environment prompted for other PPN management strategies, which involve using fewer chemicals, or bioformulation-based products.

2.2 Organic Nematode Management Due to the present impact of anthropic activities, safe solutions are needed to protect the environment and the living organisms. However, it is challenging to find the best way to solve some significant issues such as pest control. Vegetable and fruit crops are attacked by many pathogens, including fungi, bacteria, viruses, and nematodes  that are the primary factor limiting economically essential crops (Abd-­ Elgawad, 2016; Dutta et al., 2019). Present management of PPN through synthetic nematicides (where allowed by national regulations) is based on products containing active ingredients (a.i.) such as carbamate, aldicarb, oxamyl, etc. The side effects of nematicides, however, are so problematic that they disrupt the

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environment and indirectly affect the community. Nematicides work as chemicals killing PPN by diffusion in the soil or may be used as a spray against foliar nematodes such as Aphelenchoides spp. (Wright, 1981; Oka, 2010). Several alternative methods can be applied to manage PPN, such as physical techniques, crop rotation, resistant cultivars, and of course, biological methods (Dutta et al., 2019). Among all the explored methods for PPN management, biological processes appear as a suitable route for management because it is safer for our lives and the environment (Abd-Elgawad, 2016). Organic nematicides include products of natural origin or in the form of waste, applied to kill PPN effectively. Currently, the use of organic nematicides is successfully employed and gets significant attention in PPN management. Organic amendments have been used for a long time to manage PPNs. The swift reduction observed after their application in PPN populations is due to the release of toxic compounds from the decomposing materials (McSorley, 2011). Various organic nematicides, whether derived from plant parts or agricultural waste, are utilized to control PPNs. These strategies appear useful when exploring new eco-friendly tools for plant nutrition and pest management (Ntalli et  al., 2020). Organic amendments such as oil cake, green manure, agro-industrial wastes, sawdust, etc., showed a potential to reduce PPN populations.

2.2.1 Organic Nematicide Sources Plant-parasitic nematodes play a crucial role in crop field deterioration, which tremendously affects economically essential vegetables (Forghani & Hajihassani, 2020). It is, hence, necessary to manage these pathogens in a straightforward and useful way, harmless to the environment. Many studies successfully tested the potential of an organic amendment to reduce PPN severity. If we focus on our surroundings, we will often find many natural sources that are in the form of waste or grow naturally, that can be utilized to manage PPNs (Fig. 2.1). These plant-based natural products are getting more priority owing to their easily degradable, cheaper, and eco-friendly nature (Ansari et al., 2020).

2.2.2 Plants and Their By-Products Several weeds and crop wastes may possess one or more biochemical mechanisms inhibiting the functional capability of PPN (Akhtar & Malik, 2000). Many plant extracts have been found to inhibit the enzyme acetylcholinesterase in nematodes, reducing their population density. Another possible mechanism of PPN management is represented by the essential oils and plant extracts which degrade the cell membrane of nematodes and affect their permeability (Kayani et  al., 2012). Konstantopoulou et  al. (1992) also suggested that plant extracts can degrade or

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Fig. 2.1  Type of organic raw products suitable to manage PPN

denature various proteins and enzymes, interfering in the respiratory chain electron flow or ADP phosphorylation, or inhibit some related enzymes. Sukul (1992) reported that about 57 botanical families contain nematicidal compounds. The most preferred plant with an innate nematicidal activity is Azadirachta indica (Akhtar & Malik, 2000). The details of some organic nematicides are given in Table 2.1. The nematicidal potential of essential oils of various botanicals from the Asteraceae family were evaluated in in-vitro and pot experiments by Pérez et al. (2003). These authors used flowers, seeds, and roots parts of different plants viz., Chrysanthemum coronarium, C. segetum, Calendula maritima, C. officinalis, and C. suffruticosa against M. artiellia. The essential oils derived from the above plants showed a significant effect on the nematode eggs hatching. Nematode development inhibition and direct interference by C. coronarium reduced infection through a nematicidal activities that promoted the growth of chickpea. Mohan (2011) compared organic nematicides such as press mud, neem cake, neem bark, peanut oil cake, and cotton seed oil cake with inorganic nematicides like carbofuran, phorate, and aldicarbto manage Saccharum sp. PPN. The effect of the organic amendment did not reduce the nematode population compared to the inorganic control. However, the organic amendment were much safer from the environmental point of view. The essential oil of several herbs and medicinal plants tested vs PPN-parasitized plants displayed a nematicidal effect (Oka et al., 2000; Park et al., 2005). A nematicidal activity of six different weed extracts (Argemon maxicana, Achyranthes aspera, R. communis, Acalypha indica, Parthenium hysterophorus, and Trianthema portulacastrum) was observed in-vitro vs M. incognita (Khan et  al., 2017). Nematicidal activity of A. aspera, Colocasia esculenta, Monstera diliosa, Tinospora cardifolia, and Abutilon indicum was checked on M. incognita parasitized eggplant. Data showed that the above-cited plants could reduce the nematode populations (Tariq et al., 2018).

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Table 2.1  Plant-derived products and their nematicidal compounds against nematodes

Plant sources A. indica

Crysanthemum coronarium, C. segetum, Calendula maritime, C. officinalis, C suffruticosa Tagetes species (T. patula, T. erecta)

Part used/ Products PPN Leaf powder, All PPNs extract Seeds, roots and flowers,

Meloidogyne artiellia

Foliage and flowers

Pratylenchus, M. incognita

Extract and powder of whole part

Ditylenchus dipsaci and M. incognita

Artemisia dracunculus, A. verlotorum, A. absinthium Inula viscose

Leaf powder All PPNs

Palestinian arum

Leaf powder All PPNs

Canavalia ensiformis

Plant extract All PPNs

Ricinus communis

Plant extract M. incognita

Paeonia suffructicosa Cinnamomum cassia

Root extract

Soil nematodes

Soil nematodes Stem bark powder and extract Plant extract Tylenchulus Brassica species semipenetrans, such as B. napus, B. M. incognita juncea, B. oleracea, B. rapa

Nematicidal compounds and effect on nematodes References Sesquiterpenes Rjoub (2019) and lactones Oka et al. (2007a) Pérez et al. Essential oils nematicidal effect on (2003) nematode biology and eggs hatching

Polythienyls (α-terthienyl), myristic and dodecanoic acid effectively reduced nematode numbers and root gall index Flavanoids compounds of extract reduced nematode numbers Sesquiterpenic acid reduced root galling Presence of calcium oxalate, minerals, and vitamins inhibited nematode multiplication Concanavalin changed nematode host-finding behavior Lectin (ricin) reduced the M. incognita movement Essential oils nematicidal activity Essential oils nematicidal activity Organic cyanides, ionic isothiocyanate, isothiocyanate act as nematicides on nematodes

Gommers and Bakker (1988), Topp et al. (1998), Ploeg (2000), and Debprasad et al. (2000) Timchenko and Maiko (1989) and Dias et al. (2000) Oka et al. (2001) Rjoub (2019)

Marban-­ Mendoza et al. (1987) Rich et al. (1989)

Oka et al. (2001) Oka et al. (2001)

Zasada and Ferris (2003) and Lazzeri et al. (2004) (continued)

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Table 2.1 (continued)

Plant sources Sudangrass and sorghum-­ sudangrass hybrids

Manihot esculenta

Part used/ Products Whole plant degradation

PPN M. hapla

Root

PPNs

Sinapis alba, Eruca Extract sativa, Raphanus sativus

PPNs

Citrus sinensis

Orange peel meal

Cannabis sativa

Dried leaves Meloidogyne powder spp.

Meloidogyne spp.

Nematicidal compounds and effect on nematodes Cyanogenic glycoside (Dhurrin) and hydrogen cynide showed nematicidal activity Cyanogenic compound limanarin control nematode Organic cyanides, ionic isothiocyanate, isothiocyanate, and nitriles production effectively manage nematode population Control of nematode by releasing chemical compounds Active ingredients inhibit nematode

References Widmer and Abawi (2000, 2002)

Sena et al. (1982)

Soheili and Saeedizadeh (2017) and Wang and Sipes (2000)

Renčo and Kovácik (2015) Renčo and Kovácik (2015)

Much research has been done using a broad range of botanicals, which showed adequate control of PPN. Similarly, one more study was carried out by Khan et al. (2019) in which the authors tested different plants viz., Leucas cephalotes, Coccinia grandis, Commelina benghalensis, Phyllanthus amarus, and Trianthema portulacastrum on M. incognita attacking carrot. The extracts from leaves were applied at different concentrations (1000, 2000, 3000, 4000, and 5000 ppm). The result showed that the 5000  ppm concentration was the most lethal to nematodes. Zaidat et  al. (2020), investigated in-vitro and in pots, the nematicidal properties of aqueous and methanolic extracts of Ricinus communis, Sinapis arvensis, Raphanus raphanistrum, Taxus baccata, and Peganum harmala. A reduction in the hatching of the second stage of juvenile (J2) and a higher mortality were observed for M. incognita. Ntalli et al. (2020) examined the nematicidal effect of plant products and organic wastes derived from plants, which all reduced PPN numbers. Commercial seed oils of certain medicinal plants were also tested in vitro for a nematicidal property reducing hatching and increasing mortality of M. incognita (Refaat et  al., 2020). Management of M. javanica on Cicer arietinum was also obtained through organic amendments (neem cake and seed powder, vermicompost), which proved to be eco-­ friendly (Singh et al., 2020). Further data on the effect of different plant extracts on the development of various PPNs are given in Table 2.1.

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2.2.3 Organic Manures The advantages of organic waste (animal and plant residues) are remarkable for reducing nematode numbers. The application of organic manure has been shown to improve soil structure, supplying required plant nutrients, and promoting the plants growth (Agbede et al., 2014, 2017; Kaya, 2018). Organic farming is used as a key element to curtail PPN effectively. According to Atandi et al. (2017), organic farming (that received a formulation of compost, Tithonia diversifolia, and neem cake) was comparatively more effective against PPNs than conventional farming (that received fertilizers and nematicides). In a study comparing organic and inorganic fertilizers on the soil microbiota, soil columns with rye grass under greenhouse conditions were fertilized with mineral fertilizers, organic cattle slurry, or urea (P and S-free control). The rye grass rhizosphere of the slurry application showed significantly greater abundance of bacterial feeding nematodes, mycorrhizal proliferation, phosphate, sulphonate-utilizing and heterotrophic bacteria (Ikoyi et al., 2020). This study concluded that organic fertilization improved soil nematodes and microbe effects, promoting sustainable plant growth (Ikoyi et al., 2020). The potential of poultry amendment to soil against PPNs was tested through several experiments on vegetable crops (Akhtar & Malik, 2000). However, the organic amendment or compost promoted the functional properties of the biological antagonist of nematodes only if the critical products were maintained for an extended period (McSorley & Gallaher, 1995). Use of chitin as soil amendments lowered population of nematodes by releasing NH3 when decomposing in soil, activating chitinolytic microorganisms that degraded nematode eggs (Culbreath et al., 1986; Spiegel et al., 1987, 1988). Rivera and Aballay (2008) reported that M. ethiopica was managed on an infected potted vine plant (Vitis vinifera L. var. Chardonnay) using five organic soil amendments. In this study, two immature composts (one prepared from dried tea residues, broiler litter, and grapes pomace, whereas the other prepared from tea residues, rachis, and grapes pomace) were used. All the amendments were applied covering the substrate in 5 L pots in the early spring, resulting in the suppression of the nematode reproductive index (Rivera & Aballay, 2008). The activities of P. penetrans and H. glycines ceased after application of sludge processed anaerobically and hog manure (in liquid form), due to the presence of volatile fatty acids (Min et al., 2007). Mahran et al. (2008) reported that valeric acid was the most harmful to P. penetrans among all tested fatty acids, while isobutyric acid was the least effective. Organic manure is rich in many compounds, especially phenolics and nitrogen (Hassan et al., 2010; Renčo & Kováčik, 2012). Khan et al. (2018) conducted a pot study with chopped leaves of different weeds (viz., Ipomea carnea, Eichornia crassipes, Nicotiana plumbaginifolia, Ageratum conyzoides, and Trianthema portulacastrum) which reduced M. incognita numbers due to the presence of chemicals with nematicidal properties. Similarly, Asif et al. (2017c) tested six different botanicals (Solanum xanthocarpum, Achyranthes aspera, Oxalis stricta, Amarenthus spinosus, Cassia tora, and Ranunculus pensylvanicus) as

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Table 2.2  Effect of various organic manures on the multiplication of PPNs

Manure sources Poultry manure

PPN M. incognita, M. javanica, Helicotylenchus multicinctus

Farmyard manure

M. incognita

Sheep, cattle, and horse manure Cow dung and their urine

M. incognita

Chicken manure

M. incognita

Farmyard manure (mixture of cattle, sheep, and goat dung) and poultry manure

PPNs

M. incognita

Nematicidal compounds & their effect on nematodes Reduction in soil populations, galling, and eggs production, decreased severity of attack Prevented disease severity and complexity Mortality of nematode increases Reduction in nematode populationd and minimized reproduction Decreased nematode infestation level Control of PPNs

References Chindo and Khan (1990), Sundararaju et al. (2002), Kerkeni et al. (2007), Stirling et al. (2012), and Akhtar and Malik (2000) Khan (2003) and Goswami et al. (2007) Kerkeni et al. (2007) Abubakar et al. (2004)

López-Pérez et al. (2005) Rjoub (2019)

organic amendments to inhibit hatching and mortality of M. incognita. Different studies related to the application of manures against various PPNs are given in Table 2.2.

2.2.4 Agro-Industrial Wastes Agricultural wastes are non-product residues obtained from plants by the processing of raw agricultural products such as crop residues, fruits, vegetables, and from animals such as meat, poultry, and dairy products. Their content in organic matter can benefit organic agricultural farming (Obi et al., 2016; Banga & Kumar, 2019; Pandey & Dwivedi, 2020). The soil amendment of agro-industrial wastes has been one of the best methods for managing nematodes and other pests for centuries. They may cost less than other amendments, and easily degrade in fields in the form of compost, improving soil physicochemical and biological characteristics. Many studies illustrate the suppression of nematodes by microorganisms after application of animal manure (Kaplan & Noe, 1993; Oka, 2010). On average, well-formed farmyard manure (FYM) includes 0.2% P2O5, 0.5% K2O, and 0.5% nitrogen. Application of this type of FYM significantly diminished the root-knot nematode population in balsam, Impatiens balsamina (Khan, 2003).

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Using agro-industrial wastes such as silver skin (CS) and brewers spent grain (BSG) in pots lowered the population of M. incognita. During the experiment, chlorophyll content and the number of reactive oxygen species (ROS) produced by tomato plants were calculated. Compared to the untreated control, the CS extract dramatically reduced the nematode population, whereas no effect was found for the BSG extract on the final population. The highest ROS levels were triggered by CS 100%, BSG 50 and 100%, ten days after the initial treatment, which significantly impacted tomato plant growth compared to untreated controls (Thligene et al., 2019). Soil amendment with agricultural wastes viz. wheat straw, decaffeinated tea, rice husk, pigeon pea pod residue, and black gram pod residue applied at the rate of 30 and 60 g per pot significantly reduced the infestation caused by M. incognita and improved the yield of Solanum melongena (Tariq et al., 2018). Faruk (2019) showed increased onion growth and reduced gall development on roots after incorporating rice bran, mustard oil cake, and sawdust in the soil three weeks before transplanting seedlings. The addition of three commercial formulations such as Medicago sativa dry biomass, Beta vulgaris crushed to pulp, and defatted seed meal of Brassica carinata was compared on potted and field tomato cv. Regina, infested by M. incognita. These formulations were utilized in the pot and field soil at 10, 20, 30, or 40 g/ kg and 20 and 40 t/ha, respectively (D’Addabbo et al., 2020). Similarly, Osei et al. (2011) found that extracts of five organic wastes viz. cocoa bean testa, citrus waste, oil palm bunch waste, compost, and poultry manure had inhibitory effects on egg hatching of M. incognita, in an in-vitro study. Various agricultural wastes such as orange bagasse, powdered bean hulls, and soybean hulls applied at doses of 0 (control), 2, 4, 6, and 8 t/ha showed 55–100% reduction in the PPN population density (Brito et al., 2020). Effects of different agro-industrial wastes on the multiplication of various PPNs have been summarized in Table 2.3.

Table 2.3  Effect of agro-industrial wastes on the PPN development Agro-industrial wastes. Silver skin (CS), brewers’ spent grain (BSG) Black gram pod residue, decaffeinated tea, pigeon pea pod residue, rice husk, wheat straw Rice bran, mustard oil cake, sawdust

Plant studied Lycopersicum esculentum Abelmoschus esculentus

Dry biomass of Medicago sativa, pressed pulp of Beta vulgaris, defatted seed meal of Brassica carinata Orange bagasse, powdered bean hulls, soybean hulls

Lycopersicum esculentum

Allium cepa

Lycopersicum esculentum

Nematode Studied M. incognita M. incognita

Reference Thligene et al. (2019) Tariq et al. (2018)

Meloidogyne spp. M. incognita

Faruk (2019)

PPN

Brito et al. (2020)

D’Addabbo et al. (2020)

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2.2.5 Oil Cakes The incorporation of organic waste in soil provides important ingredients such as nitrogen, phosphorus, and potash (NPK). Oil cakes are by-products obtained after oil extraction from seeds. They possess nematicidal properties and therefore are used to prevent the damage caused by PPN in several crops. When soil is amended with different types of oil cakes, these products decompose in 15–20 days. After decomposition they release various compounds that improve plant growth and soil fertility and inhibit nematode populations growth (Ramachandran et  al., 2007). Several antimicrobial by-products (for example, organic acids, phenols, hydrogen sulfide, tannins, and nitrogenous compounds) are released during the decomposition of organic additives or synthesized by the breakdown of microorganisms (Rodriguez-Kabana et al., 1995). Various studies suggested that the different types of oil cake amendments may effectively control PPN in different crops (Rehman et al., 2011; Dourado et al., 2013; Patra, 2018; Baheti et al., 2019). The application of mahua and neem oil cake at 100 g/m2 reduced the number of galls in the root system of tomato attacked M. incognita (Patra, 2018). The efficacy of various oil cakes such as Gossypium hirsutum (cotton), B. campestris (mustard), and A. indica (neem) reduced the M. incognita nematode infestation (Rehman et al., 2011). The extracts of camellia seed cake showed a strong nematicidal activity against M. javanica at low concentrations (2 g/L) (Yang et al., 2015). The raw application of different oil cakes viz., karanj (Pongamia pinnata), mustard (B. campestris), and neem (A. indica) at 2, 4, and 6 q/ha reduced the number of M. incognita induced root galls in okra (Baheti et al., 2019). Resha and Rani (2015) showed that the extracts, dry leaves powder, and seeds of neem had a nematicidal activity against M. incognita by inhibiting egg hatching and causing the highest J2 mortality. Kankam et al. (2014) showed the effectiveness of oil cakes from shea nut (Vitelleria paradoxa), palm kernel (Elaes guineensis), and Indian almond (Terminalia catappa) against M. incognita attacking cowpea. The application of eight oil cakes, i.e. sunflower, linseed, cotton, sesamum, groundnut, neem, mahua, and mustard, significantly reduced the number of galls in the fibrous root of rice caused by the rice root-knot nematode, M. graminicola (Kumar et  al., 2018). Neem oil also immobilized in vitro the J2 of M. incognita (Dourado et  al., 2013). Further oil cakes viz. groundnut, neem, sunflower, and mahua cake, were very effective in reducing nematode population and increasing root nodulation in lentils (Shankar et al., 2016). Chandrawat et al. (2020) reported efficacy of oil cakes viz. castor, mahua, karanj, and mustard (at the rate of 2.5 and 5.0 q/ha and neem cake at the rate of 5.0 q/ha) against M. incognita. The enzymatic activity assayed in tomato roots revealed that the application of oilcake increased the level of defense enzymes, with time exposure. Among all, the application of mustard cake at 5.0 q/ha showed highest enzymatic activity. The soil treatments with commercial biostimulants (neem seed cake and sesame oil) suppress M. incognita multiplication and limited the nematode impact on tomato (Laquale et  al., 2018). The effects of different oil cakes on the development of various PPNs are summarized in Table 2.4.

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Table 2.4  Effect of oil cakes application on PPN multiplication and development Oil cakes. Castor, mahua, karanj, mustard Cotton, neem, mustard Shea nut, Indian almond, palm kernel Camellia Neem Groundnut, sunflower, mahua, neem Mahua, neem Cotton, neem Sesame, neem Karanj, mustard, neem

Nematode Plant studied Studied L. esculentum M. incognita

Reference Chandrawat et al. (2020) Rehman et al. (2011) Kankam et al. (2014)

C. arietinum V. sinensis

M. incognita M. incognita

M. paradisica A. esculentus Lens culinaris L. esculentum Oryza sativa L. esculentum A. esculentus

M. javanica Yang et al. (2015) M. incognita Resha and Rani (2015) Meloidogyne spp. Shankar et al. (2016) M. incognita M. graminicola M. incognita M. incognita

Patra (2018) Kumar et al. (2018) Laquale et al. (2018) Baheti et al. (2019)

2.2.6 Biochar Biochar is a pyrolyzed carbon-rich charred biomass made from crop residue, wood, manures, and other sources. It is a charcoal-like carbonaceous by-product of pyrolysis at relatively low temperatures under limited oxygen availability (Lee et  al., 2013). To boost productivity, conserve carbon reserves, or filter percolating soil water, “Biochars” are practiced as a soil amendment (Lehmann & Joseph, 2009). Thermal process settings (i.e., temperature, heating rate, etc.) and original feedstock composition influence the feature of biochars (Gaskin et  al., 2008; Singh et  al., 2010; Cantrell et al., 2012; Hmid et al., 2014), thus finding out the best choice for its final use (Antal & Grønli, 2003). The advantage of biochar on crop yield has also been well documented (Jeffery et al., 2011). These effects are linked with various features and capabilities (i) due to the alkaline factor, and soil pH increase; (ii) due to enhanced water retention capacity, increasing soil water regime; (iii) detoxifying many xenobiotics and allelopathic entities and (iv) increasing favorable microbes triggering plant growth and development’. Biochar received attention and interest as a natural fertilizer for its carbon sequestration, thereby increasing soil health and nematode management capability. Biochar is stable when applied to soils, can store soil carbon for several years while also providing increased water holding capacity and nutrient availability (Lal, 2008). It improves the supply of essential macro and micronutrients for plant growth, mainly in acidic soils (Chan & Xu, 2009; Major et  al., 2010). The most significant biochar concentration which could decrease the nematode growth in rice roots was 1.2% under pot conditions. In contrast, biochar exudates directly affect nematode survival, infectivity, and development. A natural product such as biochar is actually getting special attention and has acquired international recognition in sustainable disease management programs

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(Rogovska et al., 2017; Asif et al., 2017a; Ansari et al., 2020). It has been widely exploited in agriculture because of its benefits to soil and crop yield (Steiner et al., 2007). However, dichloropropene added to biochar, due to its high adsorption capacity, was found as a strong anti-nematode fumigant. Consequently, biochar-soil amendments may increase the appropriate dose of dichloropropene to manage nematodes efficiently (Graber et  al., 2011). While analyzing plants attacked by root-­ knot nematodes, the number of adult females in roots treated with biochar was somewhat lower than the untreated controls (Huang et  al., 2015). According to Ibrahim et al. (2018), biochar raised the soil pH, reducing the negative impacts of Meloidogyne spp., resulting in a decrease in galling and an increase in tomato growth and yield. Enhanced biochar concentrations reduced nematode galls on the roots but showed a poorer tomato plant performance. Zhang et al. (2013) confirmed a decline in the population of Pratylenchus and Hirschmanniella spp. in biochar-­ modified wheat fields. Fungivore nematode genera preferred a biochar-enriched habitat to obtain more microbial food sources (Atkinson et al., 2010). Still, PPN genera chose environments with low biochar concentrations to avoid microbe competition (Hominick, 1999). Soil tillage, nutrient availability, and crop productivity were greatly improved after adding biochar (Elad et  al., 2010; Lehmann et  al., 2003). Ebrahimi et al. (2016) reported that biochar undoubtedly reduced the hatching of potato cyst nematodes, as the final number of juveniles that reached and entered the roots during the growing period did not decline.

2.2.7 Chitosan Chitosan is a modified, natural carbohydrate polymer obtained through the deacetylation of chitin [poly-β-(1  →  4)-N-acetyl-D-glucosamine], derived from crustaceous shells such as shrimps and crabs, lobster, krill, and other crustaceans. Chitosan, a high non-toxic, bioactive polymer, has become a functional recognized molecule due to its pesticide effects and elicitation of defense mechanisms in plant tissues (Wilson et al., 1994; Terry & Joyce, 2004; Radwan et al., 2012). Its nematicidal activity has been demonstrated both in vitro and in vivo. Nematode inhibition is closely associated with concentration, indicating that chitosan performance is influenced by the rate at which it is applied. When treated to foliage or soil, chitin and its deacetylated derivative, chitosan, are effective against bacteria, viruses, fungi, and nematodes (El Hadrami et al., 2010). Chitosan application through irrigation increased the dry shoot and fresh root weight of tomato in M. javanica-inoculated plants, as well as the root length of plants inoculated with the beneficial, endophytic biocontrol fungus Pochonia chlamydosporia. Chitosan also increased the dry shoot mass of plants treated with both P. chlamydosporia and M. javanica. It dramatically enhanced P. chlamydosporia root colonization but had little effect on nematode numbers per plant or fungal egg parasitism (Escudero et al., 2017). When used alone or in combination with agricultural waste, chitosan diminished root-knot index and population of nematodes in soil (Asif et al., 2017b).

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Radwan et al. (2012) discovered that chitin and chitosan suppressed tomato root galls and numbers of M. incognita J2  in soil, in a dose-dependent manner. As a result, it was more effective than chitin in diminishing galls on roots and J2 in soil. The shoot and root lengths in soil modified with either chitin or chitosan significantly increased compared with untreated inoculated plants. Spiegel et al. (1986, 1987) observed that chitin-treated plants showed a reduction in the galling index of M. javanica and increased growth parameters of bean and tomato. Clandosam, a nematicide made up of crab shells and agricultural grade urea, effectively controlled different pests and pathogens (Fiola & Lalancettle, 2000).

2.3 Mechanisms and Modes of Action Numerous mechanisms have been introduced to explain the beneficial impact of organic amendments on plants. Many researchers showed that decomposing of organic additives release nematicidal compounds that induce plant tolerance against nematodes and/or activate natural enemies of nematodes (Akhtar & Malik, 2000; Oka, 2010; Ansari & Mahmood, 2019a, b). Multiple mechanisms are involved simultaneously, so it is hard to know which ones are the most relevant (Akhtar & Malik, 2000).

2.3.1 Release of Nematotoxic Materials Plant residues release organic acids and nitrogen-containing substances during their breakdown or disintegration in soil. Previous studies have established that these compounds possess nematotoxic properties (Oka, 2010; Thoden et  al., 2011). Table 2.5 shows plant species yielding nematotoxic, natural chemicals (Gommers & Bakker, 1988; Chitwood, 2002a). The action mechanism of genotoxic materials are not always easy to detect because they may act on multiple nematode sites or have combined effects. Nematotoxic compounds may inhibit acetylcholinesterase activity, resulting in the cession of nerve impulses and, eventually, nematode death (Nasr, 2015). Biofumigation involves the use of volatile poisonous chemicals, such as isothiocyanates produced from glucosinolates in Brassicaceous crops to reduce soil-borne infections (Kirkegaard et  al., 1998). Isocyanates have various effects on cells, including electron transport inhibition, enzyme deactivation, and signaling for cell apoptosis (Aguiar, 2012). Methyl bromide is poisonous to nematodes because of alkylated proteins and oxidization of the Fe2+ centers in the cytochrome, preventing the pathogen from respiring, just like isocyanates (Chitwood, 2002b). Some organic acids are highly toxic to PPN in vitro. However, soil characteristics, oxygen concentrations, redox potential, pH, microbial activity, and soil temperature are all

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Table 2.5  List of nematotoxic compounds with their source and tested nematode species Plant species Acacia gummifera Ageratum houstonianum Ailanthus altissima Aster sedifolius Brassica juncea Chenopodium ambrosioides Crotalaria

Gochnatia barrosii Lantana camara

L. camara

Limnanthes alba Medicago sativa

Melia azedarach

Peganum harmala Piper Tagetes

T. patula

Nematotoxic compounds Flavonoids

Nematode species Meloidogyne spp.

References El Allagui et al. (2007) 1,2-dehydropyrrolizidine alkaloids M. hapla Thoden et al. (2009) (E, E)-2,4-decadienal and M. javanica Caboni et al. (E)-2-decenal and furfural (2012) Saponins M. incognita Di Vito et al. (2010) Glucosinolates P. penetrans Zasada et al. (2009) (Z)-Ascaridole M. incognita Chuan et al. (2011) 1,2-Dehydropyrrolizidine M. incognita, H. Thoden et al. alkaloids schachtii, P. penetrans, (2009) Phasmarhabditis hermaphrodita Kaempherol M. exigua Dos Santos Júnior 3-O- β-D-(6″-p-coumaroy)et al. (2010) glucopyranoside 11-oxo triterpenic acid, pomolic M. incognita Srivastava et al. acid, lantanolic acid, lantoic acid, (2006) and camarin, lantacin, camarinin and Begum et al. ursolic acid (2008) p-hydroxybenzoic acid, vanillic M. javanica Shaukat et al. acid, caffeic acid, ferulic acid, (2003) quercetin glycoside and 7-glucoside 3-methoxybenzyl isothiocyanate M. hapla Zasada et al. (2012) Triterpeneglycosides of Xiphinema index, M. D’Addabbo et al. medicagenic acid incognita, G. (2011) and rostochiensis Leonetti et al. (2010) Acetic acid, butyric acid, hexanoic M. incognita Ntalli et al. acid, decanoic acid, furfural, (2010a, b) 5-hydroxymethylfurfural and furfurol Alkaloids Meloidogyne spp. El Allagui et al. (2007) Capsaicin Meloidogyne spp. Edelson et al. (2002) Polyacetylenes, polyethienyls and Meloidogyne spp. Chitwood (2002a, flavonoids b), Marahatta et al. (2010), and El Allagui et al. (2007) α-Terthienyl acid, gallic acid and H. zea Faizi et al. (2011) linoleic acid

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factors to consider, as they severely affect the nematicidal action of organic amendments (Taba et al., 2006). Neem has been most widely studied against PPN due to its pesticide potential. Limonoids are the principal ingredients responsible for the pesticide effect of neem, azadirachtin being the most important. Several neem preparations such as leaves, kernel oil, and oil cake have been examined against PPN (Akhtar, 2000). Oil cake, the de-oiled residue of seed, has been used as fertilizer and for nematode suppression (Akhtar & Mahmood, 1996; Akhtar, 1998b; Abbasi et al., 2005). Oil cakes are rich in nematotoxic chemicals like ammonia released during their decomposition and degradation in the soil (Abbasi et al., 2005). In combination, neem preparation also enhanced the efficacy of other organic products against nematodes (Oka et al., 2007b). Tagetes spp. and many Asteraceae plants were studied for their nematotoxic activity because of their nematicidal compounds, especially α-terthienyl (Faizi et al., 2011; Salehi et al., 2018). Tagetes spp. are commonly used as a rotational crop to manage PPN populations, especially Meloidogyne and Pratylenchus spp. However, soil amendment with Tagetes spp. is also effective against nematodes. The incorporation of T. patula foliage into M. incognita-infected soil decreased the nematode infestation level (Marahatta et al., 2012). Bar-Eyal et al. (2006) reported that the aqueous extract of Chrysanthemum coronarium displayed nematicidal activity against M. javanica J2 in vitro. Leaf matter from C. coronarium added to soil was effective against M. javanica. Incorporation of roots, leaves, flowers, or seeds of C. coronarium, and flowers of many other Asteraceae plants (such as Calendula officinalis, Chrysanthemum segetum, C. suffruticosa, and C. maritima) in soil displayed a significant reduction in the reproduction rate of M. artiellia (Pérez et al., 2003). Some leguminous plants such as Canavalia ensiformis generate lectins such as concanavalin-A, which may disturb the nematode behavior and host recognition (Rocha et al., 2017). Castor bean was also proposed as a rotational crop and manure to suppress the population of M. arenaria (Rodriguez-Kabana et al., 1989; Ritzinger & McSorley, 1998). Soil amended with castor bean oil cake was also found to have nematotoxic properties (Lopes et al., 2009). Different nematotoxic compounds and their effects on tested nematode species are given in Table 2.5.

2.3.2 Effect on Nematode Development 2.3.2.1 Soil Effects Organic products may alter the physical properties of soil, including pH, particle aggregation, salinity, CO2 and O2 concentrations, electrical conductivity, redox potential, and soil structure that may adversely affect nematode behaviors as well as their mortality, mobility, and hatching (Oka, 2010). Organic amendments generate NH3, which may help suppress PPN. Soil treatment with cement kiln dust caused its

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alkalization up to pH  11, which increased the mortality of M. javanica J2 (Oka et al., 2006). Habash and Al-Banna (2011) showed that high salinity of soil inhibited the reproduction and development of M. incognita and M. javanica. Soil amendment with animal manure may increase its salinity and electrical conductivity (Poudel et al., 2001), which could negatively affect nematodes. Organic amendments also change the soil water osmotic pressure. Feder’s (1960) experiment showed that soil amendment with sucrose and glucose at 1–5% (w/w) concentration increased the osmotic pressure, which caused direct nematode mortality within 24 h. Addition of granular sucrose and its solution to soil reducee the nematode infestation level in a tomato crop (Santiago et al., 2005). During the decomposition of organic matter due to the activity of microorganisms in soil the concentration of CO2 rises, and O2 declines. High CO2 concentrations and low or no O2 affect the PPN behavior as well as hatching, mobility, and infectivity (Kitazume et al., 2018; Banerjee & Hallem, 2020; Topalović et al., 2020). These changes are predicted to be extreme in amended soil covered with a plastic sheet. It is an effective technique for managing nematode and fungal diseases (Taba et al., 2006; Oka et al., 2007b; Van Bruggen et al., 2016). Organic amendments may also affect the soil structure that plays a crucial role in nematode suppression. It is well known that the soil pore size changes with the soil aggregation, and this change may affect the PPN habitat and mobility. Many researchers reported that smaller and larger soil pore sizes inhibit the movement of nematodes (Otobe et al., 2004; Fujimoto et al., 2010). A larger pore size may allow significant natural pests like collembolans, mites, turbellarians, enchytraeids, tardigrades, and predatory nematodes to migrate in the habitat and kill the PPN therein present. 2.3.2.2 C/N Ratios The addition of organic products with low C/N ratios in an acidic soil is also helpful to manage PPN and soil-borne diseases. Amendment using plant manure with C/N ratios (15–20) was considered most effective against nematodes. To promptly kill nematodes using materials with short C:N ratios, ammonia and other nitrogenous chemicals are required, often found in most compost generated from various organic materials. The most outstanding efficiency against PPN has been observed with amendments comprising C:N