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471 Pages·1993·15.206 MB·English
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Methods in Neurosciences Edited by P. Michael Conn Department of Pharmacology The University of Iowa College of Medicine Iowa City, Iowa Volume 14 Paradigms for the Study of Behavior ACADEMIC PRESS, INC. A Division of Harcourt Brace & Company San Diego New York Boston London Sydney Tokyo Toronto Front cover photograph: Displaying maternal behavior, a protective adult oversees the play of a young pup. Photograph by Deborah Conn. This book is printed on acid-free paper. @ Copyright © 1993 by ACADEMIC PRESS, INC. All Rights Reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher. Academic Press, Inc. 1250 Sixth Avenue, San Diego, California 92101 United Kingdom Edition published by Academic Press Limited 24-28 Oval Road, London NW1 7DX International Standard Serial Number: 1043-9471 International Standard Book Number: 0-12-185277-6 (Hardcover) (alk. paper) PRINTED IN THE UNITED STATES OF AMERICA 93 94 95 96 97 98 EB 9 8 7 6 5 4 3 2 1 Contributors to Volume 14 Article numbers are in parentheses following the names of contributors. Affiliations listed are current. J. H. ABRAINI (16), Biologie des Hautes Pressions, Faculté de Medicine Nord, 13326 Marseille Cedex 15, France ENRICO ALLEVA (7), Section of Behavioral Pathophysiology, Laboratorio di Fisiopatologia di Organo e di Sistema, Instituto Superiore di Sanità, 1-00161 Rome, Italy DAVID E. ANDERSON (26), Laboratory of Behavioral Sciences, National Institute on Aging, National Institutes of Health, Gerontology Research Center, Baltimore, Maryland 21224 JULIO M. AZCURRA (10), Departamento de Ciencias Biológicas, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Ciudad Un­ iversitaria, 1428 Buenos Aires, Argentina J. LEE BEVERLY (20), Department of Surgery, University Hospital, State University of New York, Health Science Center, Syracuse, New York 13210 RICHARD J. BODNAR (18), Department of Psychology and Neuropsychol- ogy, Queens College, City University of New York, Flushing, New York 11367 GEORG ANDREES BÖHME (14), Rhône-Poulenc Rorer, Centre de Recherche de Vitry-Alfortville, 94403 Vitry-sur-Seine Cedex, France JUAN L. BRUSÉS (10), Department of Physiology and Neurobiology, The University of Connecticut, Storrs, Connecticut 06269 THOMAS W. CASTONGUAY (19), Department of Human Nutrition and Food Systems, University of Maryland, College Park, Maryland 20742 MARIA ESTER CELIS (24), Department of Pharmacology, Facultad de Cien­ cias Quimicas, 5016 Cordoba, Argentina PAUL F. CHAPMAN (28), Department of Psychology and Graduate Program in Neuroscience, University of Minnesota, Minneapolis, Minnesota 55455 JAMES A. CHERRY (1), Cold Spring Harbor Laboratory, Beckman Neuro­ science Center, Cold Spring Harbor, New York 11724 IX X CONTRIBUTORS TO VOLUME 14 JOHN T. CLARK (3, 4), Department of Physiology, Meharry Medical College, Nashville, Tennessee 37208 MARGARET G. CUTLER (6), Department of Biological Sciences, Glasgow Polytechnic, Glasgow G4 OBA, Scotland, United Kingdom GABRIELA DÌAZ-VÉLIZ (11), Departamento Preclinicas, Facultad de Medi­ cina, Division Ciencias Médicas Oriente, Universidad de Chile, Santiago 9, Chile DEAN DLUZEN (5), Department of Anatomy, Northeastern Ohio Universi­ ties, College of Medicine, Rootstown, Ohio 44272 SAMUEL F. DWORKIN (21), Department of Oral Medicine, University of Washington, School of Dentistry, Seattle, Washington 98195 BERNARD T. ENGEL (26), Laboratory of Behavioral Science, National Insti­ tute on Aging, National Institutes of Health, Gerontology Research Center, Baltimore, Maryland 21224 KEVIN C. F. FONE (12), Department of Physiology and Pharmacology, Med­ ical School, Queen's Medical Centre, Nottingham University, Nottingham NG7 2UH, England JOHN R. GLEASON (20), Department of Statistics and Measurement, College of Arts and Sciences, Syracuse University, Syracuse, New York 13244 CHRISTOPHER J. GORDON (17), Neurotoxicology Division, Health Effects Research Laboratory, U.S. Environmental Protection Agency, Research Triangle Park, North Carolina 27711 JAANUS HARRO (23), Laboratory of Psychopharmacology, Institute of Gen­ eral and Molecular Pathology, Tartu University, Verski 34, EE-2400 Tartu, Estonia KENJI ICHIHARA (13), Department of Chemical Pharmacology, Faculty of Pharmaceutical Science, Meijo University, Nagoya 468, Japan, and Re­ search Laboratory, Maruko Pharmaceutical Co., Ltd., Kasugai, 486, Japan THEODORE J. KALOGERIS (19), Department of Physiology and Biophysics, Louisiana State University School of Medicine, Shreveport, Louisiana 71130 ERNEST D. KEMBLE (8), Division of Social Sciences, University of Minne­ sota-Morris, Morris, Minnesota 56267 PHYLLIS E. MANN (22), Department of Comparative Medicine, Tufts Uni­ versity School of Veterinary Medicine, North Grafton, Massachusetts 01536 CONTRIBUTORS TO VOLUME 14 XI PETER MARLER (29), Department of Zoology, Section of Animal Physiology, Animal Communication Laboratory, University of California, Davis, Davis, California 95616 MICHAEL M. MEGUID (20), Department of Surgery, University Hospital, State University of New York, Health Science Center, Syracuse, New York 13210 SERGIO MORA (11), Departamento Preclinicas, Facultad de Medicina, Divi­ sion Ciencias Médicas Oriente, Universidad de Chile, Santiago 9, Chile TOSHITAKA NABESHIMA (13), Department of Neuropsychopharmacology and Hospital Pharmacy, Nagoya University School of Medicine, Showa-ku, Nagoya 466, Japan DOUGLAS A. NELSON (29), Department of Zoology, Section of Animal Phys­ iology, Animal Communication Laboratory, University of California, Davis, Davis, California 95616 M. J. PARKES (15), Department of Physiology, Royal Free Hospital School of Medicine, London NW3 2PF, England EDWARD O. PRICE (2), Department of Animal Sciences, University of Cali­ fornia, Davis, Davis, California 95616 VICTOR D. RAMIREZ (5), Department of Physiology and Biophysics, Univer­ sity of Illinois, Urbana, Illinois 61801 ROBERTO REFINETTI (17), Department of Psychology, College of William and Mary, Williamsburg, Virginia 23187 MICHEL REIBAUD (14), Rhône-Poulenc Rorer, Centre de Recherche de Vi- try-Alfortville, 94403 Vitry-sur-Seine Cedex, France J. C. ROSTAIN (16), Biologie des Hautes Pressions, Faculté de Medicine Nord, CNRS-URA 1330, 13326 Marseille Cedex 15, France ZOLTAN SARNYAI (9), Alcohol and Drug Abuse Research Center, McLean Hospital-Harvard Medical School, Belmont, Massachusetts 02178 LONNIE L. SEARS (28), Department of Pediatrics, University of Louisville, Louisville, Kentucky 40292 LENNART SVENSSON (25), Department of Pharmacology, University of Goteborg, S-413 90 Goteborg, Sweden MARK I. TALAN (26), Laboratory of Behavioral Sciences, National Institute on Aging, National Institutes of Health, Gerontology Research Center, Bal­ timore, Maryland 21224 XÜ CONTRIBUTORS TO VOLUME 14 C. TOMEI (16), Biologie des Hautes Pressions, Faculté de Medicine Nord, 13326 Marseille Cedex 15, France ENRIQUE TORRE (24), Department of Neurosciences, University of Virginia, School of Medicine, Charlottesville, Virginia 22908 T. R. VIDYASAGAR (27), Center for Visual Science, and Division of Neuro­ sciences, John Curtin School of Medical Research, Canberra, ACT 2601, Australia LEANNE WILSON (21), Department of Oral Medicine, University of Wash­ ington, School of Dentistry, Seattle, Washington 98195 ZHONG-JIN YANG (20), Department of Surgery, University Hospital, State University of New York, Health Science Center, Syracuse, New York 13210 Behavior is the culmination of the gene expression, hgand-receptor interac­ tions, and all the biochemistry that makes an organism precisely that! The measurement of behaviors is, owing to its intrinsic complexity, exceedingly difficult, and approaches differ from one research group to another. This volume is dedicated to detailed descriptions of the means for measur­ ing a wide range of behaviors. It describes sexual, reproductive, and mater­ nal behaviors, social interactions, aggression, drug- and environment- induced behaviors, amnesia, maze tests, feeding behavior, illness behavior, exploratory behavior, grooming behavior, cardiovascular responses, analy­ sis of memory, and classical conditioning. The goal of this volume—and of others in this series—is to provide in one source a view of the contemporary techniques significant to a particular branch of neurosciences, information which will prove invaluable not only to the experienced researcher but to the student as well. Of necessity some archival material will be included, but the authors have been encouraged to present information that has not yet been published, to compare (in a way not found in other publications) different approaches to similar problems, and to provide tables that direct the reader, in a systematic fashion, to earlier literature and as an efficient means to summarize data. Flow diagrams and summary charts will guide the reader through the processes described. The nature of this series permits the presentation of methods in fine detail, revealing "tricks" and shortcuts that frequently do not appear in the litera­ ture owing to space limitations. Lengthy operating instructions for common equipment will not be included except in cases of unusual application. The contributors have been given wide latitude in nomenclature and usage since they are best able to make judgments consistent with current changes. I wish to express my appreciation to Mrs. Sue Birely for assisting in the organization and maintenance of records and to the staff of Academic Press for their efficient coordination of production. Appreciation is also expressed to the contributors, particularly for meeting their deadlines for the prompt and timely publication of this volume. P. MICHAEL CONN Xlll Methods in Neurosciences Volume 1 Gene Probes Edited by P. Michael Conn Volume 2 Cell Culture Edited by P. Michael Conn Volume 3 Quantitative and Qualitative Microscopy Edited by P. Michael Conn Volume 4 Electrophysiology and Microinjection Edited by P. Michael Conn Volume 5 Neuropeptide Technology: Gene Expression and Neuropeptide Receptors Edited by P. Michael Conn Volume 6 Neuropeptide Technology: Synthesis, Assay, Purification, and Processing Edited by P. Michael Conn Volume 7 Lesions and Transplantation Edited by P. Michael Conn Volume 8 Neurotoxins Edited by P. Michael Conn Volume 9 Gene Expression in Neural Tissues Edited by P. Michael Conn Volume 10 Computers and Computations in the Neurosciences Edited by P. Michael Conn Volume 11 Receptors: Model Systems and Specific Receptors Edited by P. Michael Conn Volume 12 Receptors: Molecular Biology, Receptor Subclasses, Localization, and Ligand Design Edited by P. Michael Conn Volume 13 Neuropeptide Analogs, Conjugates, and Fragments Edited by P. Michael Conn Volume 14 Paradigms for the Study of Behavior Edited by P. Michael Conn Volume 15 Photoreceptor Cells (in preparation) Edited by Paul A. Hargrave Volume 16 Neurobiology of Cytokines (Part A) (in preparation) Edited by Errol B. De Souza Volume 17 Neurobiology of Cytokines (Part B) (in preparation) Edited by Errol B. De Souza xv [1] Measurement of Sexual Behavior: Controls for Variables James A. Cherry Sexual behavior in many animals can be readily observed under standard laboratory conditions. Although often considered to be a complex form of behavior, mating typically consists of a series of stereotyped and well- characterized events that are relatively conserved between a wide range of species. Thus, with some patience and a reasonably small budget, the study of sexual behavior is within the realm of virtually any neuroscientist. This chapter outlines basic considerations for setting up the proper conditions for observing sexual behavior and then discusses possible methods of scoring and analysis. Finally, a standard procedure for observing behavior in the laboratory is described. To generalize for the largest audience, discussion focuses primarily on rodent sexual behavior. Factors Influencing Sexual Behavior and Its Measurement It is a simple matter to place adult male and female rats together and be relatively assured that behaviors of a sexual nature will occur. In fact, mating can proceed in many species in the laboratory under most conditions without apparent distraction. However, the ease with which one observes such behavior can obscure the fact that a number of important factors may contribute to both the quality and quantity of behaviors that are ultimately displayed. The strain, age, and sexual experience of animals all affect sexual performance and must be standardized. Husbandry prac­ tices that include photoperiod and housing or social environment may be equally important. Conditions of testing, including the size and complexity of the test arena, presence of odors, hormonal condition of subjects, and the time of day, will also affect performance. Fortunately, in most cases there are no "correct" conditions or factors that must be used. What is important, however, is maintaining tight control over whatever conditions are decided on. This is the best way to reduce the variability inherent in behavioral data as well as to prevent undesired influences from confound­ ing the experimental treatments. Methods in Neurosciences, Volume 14 Copyright © 1993 by Academic Press, Inc. All rights of reproduction in any form reserved. 3 4 I SEXUAL AND REPRODUCTIVE BEHAVIOR Species/Strain The question of what species or strain to examine is usually dictated by an investigator's general past usage. However, in cases where this choice can be made, there are several factors that need to be considered. For laboratory- bred rodents, strains can differ greatly with respect to reliable performance during staged matings, as well as in absolute behavioral measures such as mount frequency and ejaculation latency (1-3). If pregnancy and measures of fertility are part of the experimental design, one should be aware that fecundity is well known to vary between different strains of rats and mice. A thorough understanding of the mating cycle of a species will also be required in order to run controlled behavioral studies in the laboratory. For most standard laboratory rodents—rats, mice, hamsters—ovulation and periods of sexual receptivity occur spontaneously in regular 4- or 5-day cycles. However, for "reflex" ovulating species, including voles, rabbits, cats, and ferrets, ovulation is induced by sensory and mechanical stimulation provided by the male. In ferrets, as with many mammalian species, onset of reproductive activity occurs in response to increasing day length (see below). Unlike the spontaneously ovulating females that exhibit reoccurring cycles of receptivity, female ferrets, once in estrus, may remain sexually receptive for weeks if mating with a male does not occur. These species differences thus require modified procedures for the preparation and maintenance of reproductively active animals (see section on Stimulus Animals). Age Rodents mature quickly. Hamsters, rats, and mice are capable of reproducing anywhere from 7 to 10 weeks of age if allowed to mature naturally, but they can display sexual behaviors much earlier if provided with exogenous steroids (4, 5). Peak sexual performance in rodents can usually be observed for at least 6 months, and of course much longer for species with long life spans. Conversely, decline in reproductive output, for a variety of known and unknown reasons, occurs with age (6, 7). Sexual Experience Copulation is usually not as polished in first-time maters at it is after experi­ ence has been acquired. In hamsters, the behavior of sexually inexperienced males is often characterized by incorrect orientation toward the female,

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