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Theory and Practice of Immunocontraception in Wild Mammals Author(S): Lisa I

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Theory and Practice of Immunocontraception in Wild Mammals Author(S): Lisa I Theory and Practice of Immunocontraception in Wild Mammals Author(s): Lisa I. Muller, Robert J. Warren, Donald L. Evans Source: Wildlife Society Bulletin, Vol. 25, No. 2, Deer Overabundance (Summer, 1997), pp. 504- 514 Published by: Allen Press Stable URL: http://www.jstor.org/stable/3783483 Accessed: 05/05/2010 12:26 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=acg. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. Allen Press is collaborating with JSTOR to digitize, preserve and extend access to Wildlife Society Bulletin. http://www.jstor.org 504 IMMUNOCONTRACEPTIONIN WILDMAMMALS Theory and practice of immunocontraceptionin wild mammals Lisa I. Muller, Robert J. Warren, and Donald L. Evans Abstract Immunocontraceptionhas been proposed as a technique for managing wildlife popula- tions in urbanand suburbansettings where traditional,lethal control methods may not be publicly acceptable. Immunocontraceptionuses an animal's own immune response to disrupt reproductivefunction. Proteinsof eggs, sperm, fertilized eggs, and reproductive hormones have variously been proposed for use in developing a vaccine for fertilitycon- trol. The most widely tested immunocontraceptivevaccine for wild species is based on developing antibodies to the zona pellucida (ZP), which surroundsthe mammalianegg cell. This vaccine has successfully caused infertilityin some individual animals, but re- quires multiple treatments. Enhancementof immune response and efficiency of vaccine delivery will be necessary before this type of management strategy can be applied to wildlife control at the population level. Contraceptivetreatment may alter the health and behavior of wildlife populations and thereforemust be monitoredclosely. Keywords contraceptives, ELISA,enzyme-linked immunoabsorbentassay, Equuscaballus, immuno- contraceptives, immunosterilization,Odocoileus virginianus,population management Increasing attention is being focused on fertility American Association of Wildlife Veterinarians (re- control as a possible technique for controlling viewed by Nettles 1997). wildlife populations in urban and suburban areas. In 1991, Turner and Kirkpatrickreviewed the most In many of these areas, wildlife populations are be- recent developments in wildlife immunocontracep- coming overabundant. However, in such settings, tion. Since that time, a number of advances have traditional hunting or lethal control programs may taken place in this area of research. It is timely now not be publicly acceptable. A demand has arisen for to present an updated review. Our objectives are to contraceptive management techniques (e.g., "Bat- provide a brief review of the immunological theory tling animal overpopulation-deer herds at heart of and biology underlying immunocontraception. Many park conflict" USA TODAY,25 Mar 1993), but fertil- wildlife managers may not be interested in acquiring ity control is not a panacea for population manage- a thorough knowledge of immunology; however, ment. First, effective contraception must be there are many important details of reproductive im- achieved, and delivery of the contraceptive must be munology that are directly related to the practical practical and cost effective. There also are many an- treatment of wildlife species with immunocontracep- imal health questions that must be resolved before tives. These details often are not of concern to re- the use of contraceptives for population control is productive specialists that work with either domestic considered (Nettles 1997). These concerns were species or human subjects. Therefore, we will re- also voiced in a proposed resolution on the use of view the relevant concepts in both immunology and fertility control in free-ranging wildlife by the wildlife management when considering immunology- Wildlife Disease Association (1994) and by the based fertility control for wildlife management. Addressfor Lisa1. Muller: Department of Agricultureand NaturalResources, Delaware State University, Dover, DE 19901-2277, USA. Addressfor RobertJ. Warren:Daniel B. WarnellSchool of ForestResources, University of Georgia,Athens, GA 30602, USA. Address for Donald L. Evans:Department of MedicalMicrobiology, College of VeterinaryMedicine, University of Georgia,Athens, GA 30602, USA. WildlifeSociety Bulletin 1997, 25(2):504-514 Peer refereed Immunocontraceptionin wild mammals * Mulleret al. 505 Wildlife biologists in the future may become more in- tration through the ZP and then fusion with the egg volved in the use of contraceptives for population plasma membrane (reviewed in Yanagimachi 1994). management than they have in the past (Garrott Warren should 1995, 1995). Biologists understand A brief review of the immunocontraceptive techniques and the advan- immunology tages or disadvantages of contraceptive fertility con- relative to immunocontraception trol in order to make informed recommendations for wildlife population management. B-cell versus T-cell immune responses We review the concepts in reproductive biology When a foreign protein (i.e., an antigen) enters the and immunology necessary to understand how po- body of an animal, the immune system undergoes a tential vaccines could cause infertility. Then, we re- complex series of events to protect the animal. The view selected reproductive proteins that have been immune response consists of the humoral, or anti- tested as immunocontraceptives in wildlife. This re- body response, and the cell-mediated response. view is not designed to be exhaustive, but rather, to These responses are carried out by lymphocytes. facilitate understanding of how immunocontracep- The 2 types of lymphocytes responsible for the divi- tives have been applied to wildlife species. Many sion of labor are B cells and T cells. The B cells pro- other proteins are being isolated and tested for use as duce antibodies responsible for the humoral re- immunocontraceptive vaccines, all of which rely on sponse. The T cells are either involved in target-cell an animal's own immune system to disrupt reproduc- death (cytotoxic T cells) or provide help for both hu- tive function. Immunological concepts also are im- moral and cell-mediated immune responses (T helper portant for understanding vaccine delivery tech- cells). Activated T helper cells release factors that af- niques. Finally, we consider selected topics in popu- fect lymphocyte growth and differentiation. A subset lation health and behavior in regard to contraceptive of T helper cells is involved in delayed-type hyper- management techniques. sensitivity reaction, which includes granulomatous inflammation. A brief review of reproductive Factors affecting the immune response biology relative to from reproductive proteins immunocontraception When the body encounters foreign pathogenic proteins (antigens), generally there is a response Many elements combine to bring about successful from the immune system. Most reproductive protein reproduction in mammals, including precise coordi- targets for immunocontraception are proteins nor- nation of hormones, gamete production, fusion of mally encountered by the immune system; therefore, oocyte (egg) with sperm, and maintenance of preg- the immune response against these "self-antigens" nancy. The hypothalamic-pituitary-gonadal axis may be weak (Alexander and Bialy 1994). The ZP is a controls sperm and egg production. Successful fertil- normal protein component of the ovary, and there- ization occurs when the sperm is deposited, travels fore is a "self-antigen." up the female reproductive tract, and finally fuses Sperm may introduce foreign proteins in the fe- with the egg (reviewed in Yanagimachi 1994). male. However, the system is adapted to allow The mammalian egg is surrounded by 3 layers: the sperm entry to the female reproductive tract for fer- vitelline membrane, the ZP (a matrix of protein with tilization without generating an immune system at- attached carbohydrate groups), and a cumulus cell tack. layer (mass of loosely packed cells). Ejaculated Antibody response in the female genital tract may sperm, coated with seminal proteins undergo many be under hormonal influence (Wira and Sandoe 1977, changes before passing through the layers surround- Murdoch et al. 1982, Wira and Sandoe 1987). A drop ing the egg. Sperm membrane structure and protein in immune response in the cervical mucus at the time composition change (i.e., capacitation) as they travel of ovulation probably improves conditions for sperm through the female's reproductive tract. Seminal pro- penetration. Additionally, Parrand Parr(1990) found
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