Chemical Signals in Vertebrates 11 Jane L. Hurst, Robert J. Beynon, S. Craig Roberts and Tristram D. Wyatt Editors Chemical Signals in Vertebrates 11 Jane L. Hurst S. Craig Roberts Department of Veterinary Preclinical Science School of Biological Sciences University of Liverpool, Leahurst University of Liverpool, Neston CH64 7TE, UK Crown Street, Liverpool, L69 7ZB, UK [email protected] [email protected] Robert J. Beynon Tristram D. Wyatt Department of Veterinary Preclinical Science Office of Distance and Online Learning University of Liverpool, University of Oxford Crown Street, Liverpool, L69 7ZJ, UK Oxford OX2 7DD, UK [email protected] tristram.wyatt@continuing- education.oxford.ac.uk ISBN: 978-0-387-73944-1 e-ISBN: 978-0-387-49835-5 Library of Congress Control Number: 2007934764 C 2008 Springer Science+Business Media, LLC All rights reserved. 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Printed on acid-free paper. 987654321 springer.com Preface This volume reports the proceedings of the eleventh triennial meeting of the Chemical Signals in Vertebrates International Symposium and thus, is the 30th anniversary of the informal grouping of scientists who convene to discuss their common interests in the ways in which vertebrates use chemical signals. Previ- ous meetings were held in Saratoga Springs, New York; Syracuse, New York; Sarasota, Florida; Laramie, Wyoming; Oxford, England; Philadelphia, Pennsylva- nia; Tubingen, Germany; Ithaca, New York; Krak´ow, Poland and Corvallis, Oregon. The eleventh meeting was hosted by the Faculty of Veterinary Science at the University of Liverpool, and was held in Chester, England. CSiV 11 was the latest in a well regarded series of meetings, and was attended by about 80 scientists, with nearly 120 further co-authors, all with a common interest in vertebrate chemical sig- nalling, and its role in vertebrate behaviour. The species range was, as ever, remark- able – from lion to salamander, from mouse to elephant, from salmon to human, a biodiversity matched by the range of the substances used for communication. As might be expected from such diversity, we enjoyed a broad ranging programme that included sessions on olfactory assessment, pheromone delivery, sexual selec- tion (human and animal), urinary proteins, anti-predator responses, scent organs and their function, individual recognition, species recognition, sexual development and sexual communication (the full programme can still be viewed on the CSiV website). The meeting was launched by a thoughtful and far-reaching presentation from Milos Novotny, on “Genetic and environmental control of volatile mammalian chemosignals: structural and quantitative aspects” in which he introduced a future in which sophisticated ‘omics technologies would feature more strongly. We closed with an equally thoughtful perspective from another of the leading scientists in this field, Dietland M¨uller-Schwarze. As has been a feature of previous meetings, the group was enthusiastic, very willing to make huge species jumps and also to listen and support the younger investigators who often make their first international presentation at this meeting. This collegiate spirit extended to the meeting dinner at the Blue Planet Aquarium, where several members of the group unwittingly donated their watches, courtesy of the magician ‘Magic Matt’, to offset the meeting costs! CSiV has no funds of its own (a small float is carried from one meeting to the next) and we are totally reliant on sponsorship to supplement the registration fees. We are particularly pleased v vi Preface therefore to acknowledge the support of the European Chemoreception Organisation (http://ecro.cesg.cnrs.fr/) and also, the Dean of the Faculty of Veterinary Science at the University of Liverpool (http://www.liv.ac.uk/vets/). The group is increasingly proud of its acronym (CSiV), in part at least because of the television series featuring analytical forensic chemistry that almost shares the same name! In the period up to the meeting a low key web presence was created (http://www.csiv.org) that will act as a home for the group, but there are still no membership fees, and no restrictions on attendance at the meetings. This website will be the source of information about the twelfth meeting, at Dalhousie University in Halifax, Nova Scotia, Canada from July 28th-31st, 2009, organised by Heather Schellinck and colleagues. One name in particular was noticeably absent from the attendees at CSiV 11, that of the wonderfully passionate and insightful Bets Rasmussmen, who was sadly too ill to attend the meeting and who subsequently died on September 17th, 2006. We will remember her fine abilities as a scientist who truly straddled the biochem- ical/behavioural divide, and who did much to allow us to see one another ‘across the gap’. We remember with fondness her enthusiasm, her sense of fun, and her willingness to engage in discussions about the whole field. A tribute chapter from Tom Goodwin and Bruce Shulte captures her unique and fascinating contribution to the field and we are pleased to dedicate this volume to her memory. Finally, it is a pleasure to acknowledge the very considerable ‘behind the scenes” help from all those who helped make the conference a success. We would like to thank Anne Tuson and Marg Hedges, as well as all the members of the Mammalian Behaviour and Evolution Group and the Proteomics and Functional Genomics Group at the University of Liverpool. We would also like to express our thanks to the University of Chester Conference Team for their support and efforts at the venue. Jane L. Hurst Robert J. Beynon S. Craig Roberts Tristram D. Wyatt Contents A Tribute to L.E.L. “Bets” Rasmussen (1938–2006) ................... 1 Thomas E. Goodwin and Bruce A. Schulte Part I New Directions in Semiochemistry 1 Volatile Mammalian Chemosignals: Structural and Quantitative Aspects .................................................... 13 Milos V. Novotny and Helena A. Soini 2 Use of Automated Solid Phase Dynamic Extraction (SPDE)/GC-MS and Novel Macros in the Search for African Elephant Pheromones .. 25 Thomas E. Goodwin, Patrick A. Brown, Mindy S. Eggert, Maria G. Evola, Sam J. House, R. Grant Morshedi, Margaret E. Weddell, C. Joi Chen, Stephen R. Jackson, Yves Aubut, Jeff Eggert, Bruce A. Schulte, L. E. L. Rasmussen 3 Urinary Lipocalins in Rodenta: is there a Generic Model? ......... 37 Robert J. Beynon, Jane L. Hurst, Michael J. Turton, Duncan H. L. Robertson, Stuart D. Armstrong, Sarah A. Cheetham, Deborah Simpson, Alan MacNicoll, and Richard E. Humphries 4 The Biological Function of Cauxin, a Major Urinary Protein of the Domestic Cat (Felis catus) .................................... 51 Masao Miyazaki, Tetsuro Yamashita, Hideharu Taira and Akemi Suzuki 5 Putative Pheromones of Lion Mane and Its Ultrastructure ......... 61 M. Poddar-Sarkar, A. Chakroborty, R. Bhar and R. L. Brahmachary Part II Olfactory Response and Function 6 Using Ethologically Relevant Tasks to Study Olfactory Discrimination in Rodents .................................... 71 Heather M. Schellinck, Stephen R. Price and Michael J. Wong vii viii Contents 7 Comparisons of State and Likelihood of Performing Chemosensory Event Behaviors in Two Populations of African Elephants (Loxodonta africana) ......................................... 81 Bruce A. Schulte, Kathryn R. Bagley, Matthew Groover, Helen Loizi, Christen Merte, Jordana M. Meyer, Erek Napora, Lauren Stanley, Dhaval K. Vyas, Kimberly Wollett, Thomas E. Goodwin and L.E.L. Rasmussen 8 Olfactory Communication in the Ringtailed Lemur (Lemur catta): Form and Function of Multimodal Signals ...................... 91 Christine M. Drea and Elizabeth S. Scordato 9 Olfaction in the Gorilla....................................... 103 Peter Hepper, Deborah Wells, Patrick McArdle, Dwyer Coleman, and Mark Challis 10 Ecological Validity in the Study of Human Pheromones ........... 111 Tamsin K. Saxton, Anthony C. Little and S. Craig Roberts 11 The Influence of Sexual Orientation on Human Olfactory Function . 121 Mark J.T. Sergeant, Jennifer Louie and Charles J. Wysocki Part III Recognition within Species: Individual, Sex, Group 12 MHC-Associated Chemosignals and Individual Identity ........... 131 Peter A. Brennan 13 Pregnancy Block from a Female Perspective ..................... 141 Stuart D. Becker and Jane L. Hurst 14 The Wing-Sac Odour of Male Greater Sac-Winged Bats Saccopteryx bilineata (Emballonuridae) as a Composite Trait: Seasonal and Individual Differences ....................................... 151 Barbara Caspers, Stephan Franke, and Christian C. Voigt 15 Gender Specific Expression of Volatiles in Captive Fossas (Cryptoprocta ferox) During the Mating Season ................... 161 Barbara Renate Vogler, Frank Goeritz, Thomas Bernd Hildebrandt and Martin Dehnhard 16 Do Spotted Hyena Scent Marks Code for Clan