Table Of ContentBiosynthesis in Insects
Biosynthesis in Insects 
E. David Morgan 
Chemical Ecology Group, Keele University, UK 
RSeC 
advancing the chemical sciences
ISBN 0-85404-691-7 
A catalogue record for this book is available from the British Library 
0T he Royal Society of Chemistry 2004 
All rights reserved 
Apart from any fair dealing for the purpose of research or private study, or criticism or 
review as permitted under the terms of the UK Copyright, Designs and Patents Act, 1988, 
this publication may not be reproduced, stored or transmitted, in any form or by any means, 
without the prior permission in writing of The Royal Society of Chemistry, or in the case of 
reprographic reproduction only in accordance with the terms of the licences issued by the 
Copyright Licensing Agency in the UK, or in accordance with the terms of the licences 
issued by the appropriate Reproduction Rights Organization outside the UK. Enquiries 
concerning reproduction outside the terms stated here should be sent to The Royal Society 
of Chemistry at the address printed on this page, 
Published by The Royal Society of Chemistry, 
Thomas Graham House, Science Park, Milton Road, 
Cambridge CB4 OWF, UK 
Registered Charity Number 207890 
For further information see our web site at www.rsc.org 
Typeset by Refinecatch Ltd, Bungay, Suffolk, UK 
Printed and bound by TJ International Ltd., Padstow, Cornwall, UK
Preface 
“It is from  the behaviour  of  simple molecules  that  we  learn  our  most 
sign$cant  lessons. 
Sir Frederick Gowland Hopkins, 1913  
This book arose out of a course of lectures I was invited to give to young 
post-graduate  students at Universidade  Federal de Alagoas, Maceio, 
Brazil in 1999. Students working on practical aspects of insect pest con- 
trol through the use of natural pesticides, pheromones and hormones 
need to know something about the origin of these substances in nature, if 
they are to use their skills fully in their work. There is not much informa- 
tion available to them in a clear and elementary form in the review litera- 
ture or books, so I undertook to provide them with an introductory 
guide to biosynthetic pathways, with special emphasis on insects. I have 
subsequently expanded the notes prepared for them to make this small 
book, with the hope that it will prove useful and informative not only to 
them but to other students in science and technology, and perhaps attract 
more of them to this interesting and growing area of chemical ecology. 
Where biosynthetic processes differ between animals and plants or 
micro-organisms, I have concentrated on animal systems, but I have not 
confined myself strictly to the Insecta. The close interaction between 
plants and insects makes it necessary at times to look at plant substances 
and pathways, and I have, where important  or interesting  examples 
present themselves from other areas, strayed to other arthropods, and 
even man to complete the story. 
Because this is intended as a didactic work for young students, I have 
not used the apparatus of references as in a scholarly review. Experience 
has shown me that faced with many references to research literature, they 
may consult none of them. I have rather referred in the text to a few 
especially useful or interesting papers, and added a list of further reading 
at the end of each chapter. I have also included a few problems so that 
readers  can make  a self-assessment of their  grasp of what has been 
discussed. 
V
Acknowledgements 
I am grateful to my friends Neil Oldham, John Brand, Ralph Howard, 
Athula Attygalle, Graeme Jones and Desire Daloze for reading various 
parts or drafts and giving me many helpful suggestions, and to Jane 
Parker for giving me a student view on an early draft. The request for the 
original lectures came from my former student Ruth do Nascimento, to 
whom I am much indebted. I thank the British Council office in Recife, 
and CNPQ for the financial support that enabled me to give the lectures, 
and Professor Ant6nio Euzebio G. Sant’Ana for his hospitality. I am 
very grateful to Dr John Shanklin of Brookhaven National Laboratory 
and Dr Ed Cahoon of the Danforth Plant Sciences Center for kindly 
providing the diagrams of castor oil desaturase, Prof John Mann and 
Dr Rishuo Nishida for permission to use figures from their work. I am 
obliged to Dr Jonathan Banks in Australia and Dr Keith S. Brown in 
Brazil for helping me concerning the final position of aphinin research. 
Those of us laboratory-bound do not get to see a great variety of 
insects in their natural surroundings. I can happily thank various friends 
for  contributing  their  photographs  of  (biosynthetically) interesting 
insects.  Stefan0 Turillazzi, Universita di  Firenze,  for his  picture  of 
Polistes wasps, Athulla Attygalle and Maria Eisner for Epilachnis pupae, 
Steve McWilliam of rECOrd, Chester, for  Coccinella septempunctata 
adults and Jim Klaisch of the Department of Entomology, University of 
Nebraska  Lincoln for the C. septempunctata larva, Dr Mike Quinn, 
- 
Texas A&M University, Stephenville, for superb pictures of Hippodamia, 
Oncopeltus and Danaus, Tom Larsen for Scolopendra, Jens Christian 
Schou for Arctia caja, Warren E. Savary for Lytta magister, Markku 
Savela for Harpaphe haydeniana, Dr Hamish Robertson of the South 
African  Museum  for  Ceroplastes, Dr  Paul  Choate,  Department  of 
Entomology and Nematology, University of Florida for Manduca sexta 
larva, Socikte Nouvelle des Editions Boubee for permission to reproduce 
the  photo  of  a  female  silk  moth  taken  by  Jacques  Six,  Anthony 
Papadoupolos for the fire bug Pyrrhcorus apterus, the University of 
Oklahoma Veterinary School for Amblyomma americanum, and our own 
Terry Bolam for the Myrmica ant trail-following. The cover photo is one 
vii
... 
Vlll  Acknowledgements 
of the many beautiful insect photos taken by Ken Preston-Mafham of 
Premaphotos Wildlife. I have tried to find the owner of the picture of the 
cheese mite, Tyrophagus putrescentiae without success. 
I have sought to locate owners of all reproduced material not in my 
own possession. In a few cases I have been completely unsuccessful, but 
trust I have not inadvertently infringed any copyrights. Should I have 
done so I shall of course take appropriate action for any subsequent 
editions. I would also welcome comments and suggestions on this book. 
(e.d .m  [email protected] eele.a c.u k)
Description:The chemical study of insects has been growing for four decades, and with it an interest in how insects make their pheromones, hormones, defensive secretions, venoms, pigments and surface coverings. By investigating the biosynthesis of insects, one can gain a greater insight into the structure and f