prelims00.qxd 24/5/04 12:48 pm Page i NEMATODE BEHAVIOUR prelims00.qxd 24/5/04 12:48 pm Page ii prelims00.qxd 24/5/04 12:48 pm Page iii N B EMATODE EHAVIOUR Edited by Randy Gaugler and Anwar L. Bilgrami Department of Entomology Rutgers University New Brunswick New Jersey, USA CABI Publishing prelims00.qxd 24/5/04 12:48 pm Page iv CABI Publishing is a division of CAB International CABI Publishing CABI Publishing CAB International 875 Massachusetts Avenue Wallingford 7th Floor Oxfordshire OX10 8DE Cambridge, MA 02139 UK USA Tel: +44 (0)1491 832111 Tel: +1 617 395 4056 Fax: +44 (0)1491 833508 Fax: +1 617 354 6875 E-mail: [email protected] E-mail: [email protected] Web site: www.cabi-publishing.org ©CAB International 2004. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Nematode behaviour / edited by Randy Gaugler and Anwar L. Bilgrami. p. cm Includes bibliographical references and index. ISBN 0-85199-818-6 (alk. paper) 1. Nematode --Behaviour. I. Gaugler, Randy. II. Bilgrami, Anwar L., 1956- QL391.N4 N37 2004 592´.5715--dc22 2003025545 ISBN 0 85199 818 6 Typeset by MRM Graphics Ltd, Winslow, Bucks Printed and bound in the UK by Cromwell Press, Trowbridge prelims00.qxd 24/5/04 12:48 pm Page v Contents Contributors vii Dedication ix Introduction and Overview xi 1 Ecological and Behavioural Adaptations 1 Gregor W. Yeates 2 Locomotion Behaviour 25 A.H. Jay Burr and A. Forest Robinson 3 Orientation Behaviour 63 Ekaterini Riga 4 Feeding Behaviour 91 Anwar L. Bilgrami and Randy Gaugler 5 Reproductive Behaviour 127 Robin N. Huettel 6 Ageing and Developmental Behaviour 151 E.E. Lewis and E.E. Pérez 7 Osmoregulatory and Excretory Behaviour 177 Denis J. Wright v prelims00.qxd 24/5/04 12:48 pm Page vi vi Contents 8 Physiological and Biochemical Basis of Behaviour 197 Roland N. Perry and Aaron G. Maule 9 Molecular Basis for Behaviour 239 Maureen M. Barr and Jinghua Hu 10 Biotic Interactions 277 Patricia Timper and Keith G. Davies 11 Abiotic Factors 309 Mary E. Barbercheck and Larry Duncan 12 Population Dynamics 345 Brian Boag and Gregor W. Yeates 13 Survival Strategies 371 David A. Wharton Index 401 prelims00.qxd 24/5/04 12:48 pm Page vii Contributors Mary E. Barbercheck,Department of Entomology, Pennsylvania State University, University Park, PA 16802, USA (e-mail: [email protected]) Maureen M. Barr, School of Pharmacy, University of Wisconsin-Madison, 777 Highland Avenue, Madison, WI 53705, USA (e-mail: [email protected]) Anwar L. Bilgrami, Department of Entomology, Rutgers University, New Brunswick, NJ 08901-8524, USA (e-mail: [email protected], [email protected]); permanent address: Section of Nematology, Department of Zoology, Aligarh Muslim University, Aligarh 202002, India Brian Boag, Scottish Crop Research Institute, Invergowrie, Dundee, DD2 5DA, Scotland, UK (e-mail: [email protected]) A.H. Jay Burr, Department of Biological Sciences, Simon Fraser University, Burnaby, BC, V5A 1S6, Canada (e-mail: [email protected]) Keith G. Davies,Nematode Interaction Unit, Rothamsted Research, Harpenden, Hertfordshire AL5 2JQ, UK (e-mail: [email protected]) Larry Duncan, Department of Entomology and Nematology, University of Florida, IFAS, Citrus Research and Education Center, 700 Experimental Station Road, Lake Alfred, FL 33850, USA (e-mail: [email protected]fl.edu) Randy Gaugler,Department of Entomology, Rutgers University, New Brunswick, NJ 08901-8524, USA (e-mail: [email protected]) Jinghua Hu, School of Pharmacy, University of Wisconsin-Madison, 777 Highland Avenue, Madison, WI 53705, USA (e-mail: [email protected]) Robin N. Huettel, Department of Entomology and Plant Pathology, Auburn University, AL 36849, USA (e-mail: [email protected]) E.E. Lewis, Department of Entomology, Virginia Tech, Blacksburg, VA 24061- 0319, USA (e-mail: [email protected]) vii prelims00.qxd 24/5/04 12:48 pm Page viii viii Contributors Aaron G. Maule, Parasitology Research Group, School of Biology and Biochemistry, Queen’s University Belfast, Medical Biology Centre, 97 Lisburn Road, Belfast BT9 7BL, UK (e-mail: [email protected]) E.E. Pérez, Department of Entomology, Virginia Tech, Blacksburg, VA 24061- 0319, USA (e-mail: [email protected]) Roland N. Perry, Plant Pathogen Interactions Division, Rothamsted Research, Harpenden, Hertfordshire AL5 2JQ, UK (e-mail: [email protected]) Ekaterini Riga, Washington State University, 24106 North Bunn Road, Prosser, WA 99350, USA (e-mail: [email protected]) A. Forest Robinson, USDA ARS, Southern Crops Research Laboratory, 2765 F & B Road, College Station, TX 77845, USA (e-mail: [email protected]) Patricia Timper, USDA-ARS, Crop Protection and Management Research Unit, PO Box 748, Tifton, GA 31793, USA (e-mail: [email protected]) David A. Wharton, Department of Zoology, University of Otago, PO Box 56, Dunedin, New Zealand (e-mail: [email protected]) Denis J. Wright, Department of Biological Sciences, Imperial College London, Silwood Park Campus, Ascot, Berkshire SL5 7PY, UK (e-mail: [email protected]) Gregor W. Yeates,Landcare Research, Private Bag 11052, Palmerston North, New Zealand (e-mail: [email protected]) prelims00.qxd 24/5/04 12:48 pm Page ix Dedication This volume is dedicated to the memory of Neil Argo Croll (1941–1981), Professor of Parasitology at McGill University and Director of McGill International, whose untimely death prevented his witnessing the profound influence his work would have on the discipline of nematology. His legacy includes 90 research papers and seven books, including Ecology of Parasites, which was written even before completing his PhD in 1966, and the seminal Behaviour of Nematodes. In addition to the PhD, he was awarded the Doctor of Medicine degree and the presti- gious Doctor of Science degree from London University. He was particularly passionate about nem- atode behaviour and crossed over traditional boundaries to work with animal- and plant-parasitic as well as free-living marine and freshwater species. It is remarkable that one man could accomplish so much in the mere 39 years allot- ted, and we can only wonder as to ‘what might have been’ given a full lifespan. ix