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Pythium Diagnosis, Diseases and Management Editors Mahendra Rai Department of Biotechnology, SGB Amravati University Amravati, Maharashtra, India Kamel A. Abd-Elsalam Unit of Excellence in Nano-Molecular Plant Pathology Plant Pathology Research Institute, Giza, Egypt Avinash P. Ingle Department of Biotechnology, Engineering School of Lorena, University of Sao Paulo, Lorena, SP, Brazil p, A SCIENCE PUBLISHERS BOOK Cover illustration provided by Prof. Francois Barja. Designing assistance from Bengisu Sengun. Cover figures have been taken from Chapter 11 of this book. Reproduced by kind permission of the authors, Reza Cover illustration provided by Prof. Francois Barja. Designing assistance from Bengisu Sengun. Mostowfizadeh-Ghalamfarsa and Fatemeh Salmaninezhad, Department of Plant Protection, Shiraz University, Shiraz, Iran. CRC Press TCaRyClo Pr r&es Fsr ancis Group 6T0ay0l0o rB &ro kFernan Scoisu nGdr oPuaprkway NW, Suite 300 B60o0c0a RBarotoknen, F SLo 3u3n4d8 P7a-2r7k4w2ay NW, Suite 300 Boca Raton, FL 33487-2742 © 2020 by Taylor & Francis Group, LLC C© R2C02 P0r besys T isa yalno ri m&p Frriannt coifs TGaryolourp &, L FLrCancis Group, an Informa business CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works No claim to original U.S. Government works Printed on acid-free paper VPreirnstioedn oDna taec:i d20-f1r9e1e0 p0a3per Version Date: 20191003 International Standard Book Number-13: 978-0-367-25941-9(Hardback) International Standard Book Number-13: 978-0-367-25941-9 (Hardback) This book contains information obtained from authentic and highly regarded sources. Reasonable efforts hTahvies bboeoenk cmoandtae intos ipnufborlimsha trieolnia obblet adinatead afrnodm i nafuotrhmenattiioc na,n bdu ht itghhel ya ruetghaorrd eadn ds opuurbceliss.h Rere acsaonnnaobtl ea esfsfuomrtes rheasvpeo nbesiebni limtya dfoer ttoh ep vuablliidsiht yr eolfi aablll em daatetar iaalnsd o ri ntfhoer mcoantsioenq,u ebnuct etsh oef atuhtehiro ru saen. dT hpeu baulitshhoerr s caanndn optu balsissuhmeres hreasvpeo antstiebmilpittye dfo tro t threa cvea tlihdei tcyo opfy railgl hmt ahtoelrdiaerlss oorf athlle m caotnesreiaqlu reenpcreosd uocf etdh eiinr tuhsies. pTuhbeli caauttihoonr as nadn dap poulobgliiszhee tros choapvey raitgthetm hpotledde rtos itfr paceer mthies scioonpy troi gphutb hlioslhd einrs t ohfi sa lflo mrmat heraisa ln roetp breoednu coebdt aiinn tehdi.s I pf uabnlyi ccaotpioynr iagnhdt mapaotleorgiaizl eh atos ncoopt ybreiegnh ta hckonldoewrsle idf gpeedr mpliesassioen w troit peu abnldis lhe ti nu st hkins ofowr mso hwaes mnoaty breecenti foyb itna ianneyd f. uItf uarney r ceoppriynrti.ght material has not been acknowledged please write and let us know so we may rectify in any future reprint. Except as permitted under U.S. Copyright Law, no part of this book may be reprinted, reproduced, transmit- tEexdc,e oprt uatsi lpizeremd iintt eadn yu fnodremr Uby.S a. nCyo epleycritgrohnt iLca, wm, encoh paanrict aolf, tohri os tbhoeork m meaany sb, en roewp rkinnotewdn, roerp hroedreuacfetder, tirnavnesnmteidt-, itnedcl,u odri untgi lpizheodt oinc oapnyyi fnogr,m m biyc raonfyil meleinctgr,o annicd, mreeccohrdainnicga, lo, ro ri no tahneyr minefaonrms, antoiown k sntoowrang eo ro hr erreetarfiteevra iln svyesntteemd,, winictlhuoduint wg rpihttoetno cpoeprmyiinsgs,i omn ifcrroomfi ltmhein pgu, balnisdh erresc.ording, or in any information storage or retrieval system, without written permission from the publishers. For permission to photocopy or use material electronically from this work, please access www.copyright. cFoomr p (ehrtmtpi:s/s/iwonw wto.c pohpoytroigchotp.cyo omr /u) soer mcoantetraicatl tehleec Ctroopnyicraigllhyt fCrolema rtahnics ew Coerkn,t eprl,e Iansce . a(cCcCesCs) ,w 2w2w2 .cRoopsyerwigohotd. Dcormiv e(,h Dttapn:/v/ewrws, wM.cAop 0y1r9ig2h3t, .c9o7m8-/7)5 o0r- 8c4o0n0t.a CctC tChe i sC ao pnyorti-gfohrt- pCrloefairt aonrcgea nCieznattieorn, Itnhca. t( CprCoCvi)d, e2s2 l2i cRenosseews oanodd rDergiivset,r aDtiaonnv eforsr, aM vaAr i0e1ty9 2o3f ,u 9s7er8s-.7 F5o0r- 8o4rg0a0n. iCzaCtCio niss at hnaot th-faovre- pbreoefnit gorragnatnediz aa tpiohno ttohcaotp pyr loicveidnesse lbicye tnhsee Cs CanCd, are sgeipstarraattieo nsy fsotre ma v oafr pieatyym ofe nuts ehrass. Fboeer no ragrarnanizgaetdio.ns that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice:Product or corporate names may be trademarks or registered trademarks, and are used oTnralyd feomr iadrekn Ntiofitciacteio: nP raonddu ecxt polra cnoartpioonr awteit nhaomute isn mteanyt btoe itnrafrdienmgea.rks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging‑in‑Publication Data LibLriabrrya royf oCfo Cnognregsrse sCsa Ctaaltoagloinggin-ign‑-inP‑uPbulibclaictiaotnio Dn aDtaat a Names: Sengun, Ilkin Yucel, editor. TNiatmleN:e Asa:m cSeeetnisc:g Raucanii,d, I Mblkaaicnht eeYrnuidac re:a lf,,u eendddiittaoomrr..e |n Atablsd -aEnlds afloaomd, aKpapmliceal tAio.n, 1s 9/6 e9d-i teodri,tor. | Ingle, ITlkitilne : Y AAucvceientli acSs eahnc Pigd.u, bnead, cEittgoecrr .i[ a i. e: .f uEngde]a Umneinvtearlssi tayn, dE nfogoinde aeprpinligc aFtaicounlst y/ ,e Fdoiotodr, EIlnkginiTn Yeituelcerie: nlP gSy,e tDhnigeupumanr, :tE mdgiecan g[itn. eIoz.s mEisg,ie rd,] iTUsuenariksveeesyr .saintyd, mEnagniangeeemrineng tF /a ceudlittoy,r Fs:o Modahendra DEnegsci nr ieRpeatriiio,n nKg:a, BmDoeecpla aA Rr.ta mAtobennd,t -F EILzls m:a CliarRm, TC,u APrrvkeiensysa.,s [h2 0P1. 6In] g| lSee. r ies: Food biology series |D “eAs csrciipetniocen :p Buobclias hReartso bno, FokL. ”: C| IRnCc lPurdeesss b, [i2b0li1o6g]r |a pShericieasl: rFeofeorde nbcioelso agnyd series Description: Boca Raton, FL : CRC Press, [2019] | Includes bibliographical i|n “dAe sxc.ience publishers book.” | Includes bibliographical references and references and index. Iinddenexti.fiers: LCCN 2016050024| ISBN 9781498763691 (hardback : alk. paper) | IIdSBenNtI di9fei7en8rt1si:4fi L9eC8rs7C:6 LN3C7 20C07N1 (6e 20-b051o09o00k24 4: 0|a 7IlkS7.B0 p N|a Ip9Se7Br8)N14 99788706336679215 (9h4a1rd9b (ahcakr d: caolkv.e pr)a p er) | SISuBbNjeS cu9tb7s8j: e1Lc4Ct9sS8: H7L6C: 3AS7cH0e7:t oP(eby-atbhcotiueormk. :|. aF| loPko.h dpy-at-opMpeari)cthroobgieonloicg fyu. n| gBii.o reactors. CSulabsjesCicfltaiscs:as LtifiiCocnSaH:t iLo: CAnC:c Le QtCoRCb8a 2Qc.tPKe7r6.82 | 1A F.P3o49o d2P-09-1M86 2 i|c0 Dr1o9Db |Cio D l5oD7g9Cy..3 |5/ 37B9-i-o.5dr-ce-2adc3ct2o3rs . LCCla sresLicCfoic rradet ciaoovnrad:i l LaaCvbalCeil aaQtb Rlhe8t t2apt.P sh:7/t/8tlp cAsc:3n/4/.ll oc2cc0n.1g.6olo v|c /D2.g0Do1vC6/ 02550701909.320/4430--7d7c02 3 LC record available at https://lccn.loc.gov/2016050024 Visit the Taylor & Francis Web site at Vhtitspit: /t/hwew Twa.ytlaoyrl o&r aFnrdanfrcains cWise.cbo smit e at http://www.taylorandfrancis.com and the CRC Press Web site at ahnttdp t:/h/we CwRwC.c rPcrpersess Ws.ceobm si te at http://www.crcpress.com Preface The genus Pythium is one of the most important fungi of class Oomycota. It is a well-known soil- borne phytopathogenic fungus causing significant damage in agriculture, forest, nurseries, etc. and is ubiquitously found across the world. It also affects the seed germination and attacks the seedlings both at pre- and post-emergence stage. It is an unseen enemy of the root zone of various plants and is hence considered as “hidden terror” for a number of plants. The accurate diagnosis of Pythium causing root rot in plants is very important because it is often confused with root rots caused by various other fungi such as Phytophthora, Rhizoctonia, Chalara, Cylindrocladium, Fusarium and Aphanomyces. Taxonomic identification of Pythium species is also important as they vary in their host range and temperature requirements. It is proposed that Pythium root rot is difficult to control once rot has begun; therefore, its effective and eco-friendly management is a major concern. In addition, Pythium is responsible for causing infections in different animals including horses, dogs, human being, etc. and such infections are known as Pythiosis. Considering these facts, the present book is focused on all the important aspects related to Pythium biology. Broadly, this book is divided into four parts: Part I deals with an overview, host range and plant diseases, Part II presents various challenges in the taxonomy and diagnosis, and also current technological developments in detection and diagnosis of Pythium, Part III is devoted to the role of Pythium as human pathogen and Part IV is focused on the management of Pythium. The text in each chapter is supported by numerous clear, informative tables and figures. Each chapter contains relevant references of published articles, which offers a potentially large amount of primary information and further links to a nexus of data and ideas. All the chapters included in the present book have been written by the specialists, experts in the concerned topic and these chapters are highly informative and detailed. Therefore, we believe that this book will serve as a rich guide for undergraduate or graduate students of various disciplines like agriculture, plant pathology, plant physiology, mycology, molecular biology, biotechnology, and allied subjects. In addition, the book will be useful for researchers in these fields and the people working in various agro-based industries, regulatory bodies, food and agriculture organizations. The editors are highly thankful to all the contributors for their outstanding efforts to provide state-of-the-art information on the subject matter for their respective chapters. The collective efforts taken by all the authors will help to enhance and update the knowledge of the readers, particularly about Pythium and its pathogenesis in both plants and animals. We express our sincere thanks to the publisher and the authors of the chapters, whose research work have been cited in the book. We are also thankful to the entire team at CRC Press for their generous cooperation and efforts in producing this book. MR wishes to thank University Grants Commission, New Delhi for the award of BSR Faculty Fellowship. We hope that the book will be useful for all the readers to find the required information on the latest research and advances in the field of Pythium biology. Mahendra Rai Kamel A. Abd-Elsalam Avinash P. Ingle Contents Preface iii Part I: Incidence, Host-range and Diseases 1. The Genus Pythium: An Overview 3 Mahendra Rai, Kamel A. Abd-Elsalam, Avinash P. Ingle, Priti Paralikar and Pramod Ingle 2. The Genus Pythium in Three Different Continents 15 Hani Mohamed Awad Abdelzaher, Shaima Mohamed Nabil Moustafa and Hashem Al-Sheikh 3. Pythium: Diseases and Their Management 30 Patrycja Golińska and Magdalena Świecimska 4. The Genus Phytopythium 45 Abdul Mubeen Lodhi, Saleem Shahzad and Rehana Naz Syed 5. Top Three Plant Pathogenic Pythium Species 77 Amal-Asran and Kamel A. Abd-Elsalam 6. Pythium Species Associated With Die-back Apple Trees and Citrus Gummosis in Tunisia 92 Naima Boughalleb-M’Hamdi, Najwa Benfradj and Souli Mounira 7. Pathogenic and Beneficial Pythium Species in China: An Updated Review 107 Hon H. Ho and Kamel A. Abd-Elsalam 8. The Pythium Complex of the Mid-North Region of Brazil 123 Janete Barros da Silva, Douglas Henrique Trigueiro Silva, Francynara Pontes Rocha, Givanilso Cândido Leal, Helanny Márcia Ribeiro Trajano, Joseane Lustosa Machado, Laércio de Sousa Saraiva, Maria do Amparo de Moura Macêdo, Osiel César da Trindade Júnior, José de Ribamar de Sousa Rocha, Nayara Dannielle Costa de Sousa and Tamyres Lopes Rios 9. Pythium spp. on Vegetable Crops: Research Progress and Major Challenges 136 Pratibha Sharma, Prashant P. Jambhulkar, Raja, M. and Shaily Javeria 10. Host Plants and Specificity of the Genus Pythium 162 Kamel A. Abd-Elsalam and Amal-Asran Part II: Identity and Taxonomy 11. Taxonomic Challenges in the Genus Pythium 179 Reza Mostowfizadeh-Ghalamfarsa and Fatemeh Salmaninezhad vi Pythium: Diagnosis, Diseases and Management 12. Diagnosis of Pythium by Classical and Molecular Approaches 200 Shivannegowda Mahadevakumar and Kandikere Ramaiah Sridhar Part III: As a Human Pathogen 13. Pythium insidiosum – An Emerging Mammalian Pathogen 227 Erico S. Loreto, Juliana S.M. Tondolo and Janio M. Santurio Part IV: Management of Diseases Caused by Pythium 14. Damping-off Caused by Pythium Species: Disease Profile and Management 257 Mohammad Imad Khrieba 15. The Genus Pythium: Genomics and Breeding for Resistance 270 Ramadan A. Arafa, Said M. Kamel and Kamel A. Abd-Elsalam 16. Role of Phytochemicals in Plant Diseases Caused by Pythium 287 Rajendra M. Gade, Mahendra Rai, Ranjit S. Lad and Amol V. Shitole 17. Pythium aphanidermatum and Its Control Measures 299 Tahira Parveen, Mukesh Meena, Tripta Jain, Kavita Rathore, Surbhi Mehta and Kanika Sharma 18. Management of Pythium Diseases 314 Rehana Naz Syed, Abdul Mubeen Lodhi and Saleem Shahzad 19. Management of Pythium spp. by Arbuscular Mycorrhizal Fungi 344 Sarika R. Bhalerao, Pratiksha R. Gund, Sunita D. Bansod and Mahendra Rai 20. Pythium Species as Biocontrol Agents 360 Mousa Alghuthaymi, Khaled Kasem, Omar Atik and Kamel A. Abd-Elsalam Index 379 About the Editors 381 Color Section 383 CHAPTER 1 The Genus Pythium: An Overview Mahendra Rai1*, Kamel A. Abd-Elsalam2, Avinash P. Ingle3, Priti Paralikar1 and Pramod Ingle1 1 Nanobiotechnology Lab, Department of Biotechnology, SGB Amravati University, Amravati - 444602, Maharashtra, India 2 Unit of Excellence in Nano-Molecular Plant Pathology, Plant Pathology Research Institute, 9 Gamaa St., 12619 Giza, Egypt 3 Department of Biotechnology, Engineering School of Lorena, University of Sao Paulo, Estrada municipal do Campinho, sn, 12602-810 Lorena, SP, Brazil Introduction Pythium is a soil-borne pathogen which contains more than 300 species and the majority of them are plant pathogenic. Pythium is classified into 10 different clades based on morphological and genetic characteristics (Rossman et al. 2017). It is commonly found in soil, sand, various water sources and dead and decaying part of plants. It is ubiquitously distributed across the globe including America, Asia, Africa and Australia. The old taxonomic criteria proposed was confusing and there were some difficulties in the validation of various Pythium species. Pringsheim discovered the genus Pythium for the first time in 1858, and placed it in the family Saprolegniaceae (Pringsheim 1858). Thereafter, a number of attempts were made to propose different taxonomic systems for the classification of Pythium but all such systems were rejected time to time by various taxonomists (Ho 2018). According to current taxonomic system, Pythium is placed in family Pythiaceae, order Pythiales, class Oomycetes, phylum Oomycota, and kingdom Chromista (Kirk et al. 2008). As discussed earlier, Pythium spp. are mainly pathogenic to a wide variety of crop plant families and is a major problem in greenhouses and nurseries. Root rot and damping-off are the most important diseases caused by Pythium (Rai et al. 2018). It is reported that Pythium can infect some important stages of plant’s growth which mainly include infection to the seed before germination or during germination. It may attack the young seedlings before or just after emergence. The causative agents generally feed on the root system causing damping-off, which ultimately results in poor germination and spindly plants (McKellar and Nelson 2003). The infection also leads to shortened or distorted leaves, fewer tillers and smaller heads; collectively, it leads to a great loss in crop yield and economic loss (http://www.syngenta-us.com/prodrender/imagehandler.ashx?ImID=37907769 8aca-45f1-9ed2-71ad09069464&fTy=0&et=8). In addition, it is proposed that, among different species of Pythium, P. insidiosum is the most prevalent etiologic agent responsible for pythiosis in mammals (Krajaejun et al. 2018). However, among mammals, pythiosis is commonly reported in dogs, horses and humans. Apart from these, sometimes it is also observed in other animals, such as calves (Perez et al. 2005), cats (Rakich et al. 2005), sheep (Santurio et al. 2008), a bird (Pesavento et al. 2008), etc. It is also responsible for rare, non-transmissible disease generally found in tropical, subtropical and temperate regions (Mendoza *Corresponding author: [email protected]; [email protected]

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