Chemical Fertility Control and Wildlife Management

Author(s): Jay F. Kirkpatrick and John W. Turner, Jr.

Source: BioScience, Vol. 35, No. 8 (Sep., 1985), pp. 485-491
Published by: Oxford University Press on behalf of the American Institute of Biological Sciences
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 Chemical Fertility Control and Wildlife
Management

Jay F. Kirkpatrick and John W. Turner, Jr.

Chemical fertility control is a neglected yet potentially powerful wildlife management tool tion Agency before the program may
that represents an effective, inexpensive, and humane alternative to current control begin. The shortcomings of poisoning
methods. A large number of new contraceptive agents, designed primarily for use in are multiple and serious. First, the target
humans, can be administered remotely by injection to large mammals or orally to smaller animals are destroyed in a less-than-
animals. These compounds will not cause death, do not disturb social order, and can be
humane fashion, healthy animals along
manipulated to protect nontarget species.
with diseased ones. Second, population
reduction is only temporary, and each
new breeding season results in new in-
Uncontrolled or poorly controlled in- even countries. Live-trapping and relo- creases. Third, the poison kills nontarget
creases in wild and feral animal popula- cation of overpopulated species is ex- species.
tions are a problem today in many parts pensive and works only where sufficient With the notable exception of the rat,
of the world. Protection afforded certain suitable habitat exists. Poisoning over- the concept of chemical fertility control
species through refuges, as for the elk populated animals is distasteful, often as a means of controlling wild and feral
herd in Yellowstone National Park, or dangerous to humans, and notoriously populations has received surprisingly lit-
through legislation, as for feral horses nonspecific. Most poisoning programs tle attention despite a significant backlog
and burros in North America, has result- require permission from governmental of research predicting success. Thus the
ed in animal herds that exceed the land's agencies like the Environmental Protec- concept, which is not entirely new, is
carrying capacity. In other cases, popu-
lation control is needed for predatory
species (coyotes), to prevent the spread
of communicable or contagious disease
(skunks), or because of diminished habi-
tat (elephants).
Traditionally, these population in-
creases among wild and feral species
have been controlled through hunting,
trapping, relocation, and poisoning.
Controlled hunting, although successful
in certain cases, is coming under in-
creased public scrutiny. Trapping, par-
ticularly with leg-hold devices, is ex-
tremely unpopular among certain
segments of society, and legislation
against steel traps has been passed or is
pending in many states, provinces, and
Kirkpatrickis with the Departmentof Biological
Sciences, EasternMontanaState College, Billings,
MT59101;Turneris withthe Departmentof Physiol-
ogy, MedicalCollegeof Ohio, Toledo, OH 43699. ?
1985AmericanInstituteof Biological Sciences. All
rightsreserved. "Itsays here that we eat these cabbages for21 days and then we eat... "

September 1985 485

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largelyuntested, and skeptics often con-
sider the approachbizarre. The technol-
ogy associated with chemical fertility
control in humansis impressive, and its
application to wild or feral species is
fundamentally sound. The compounds
available for use in humans were first
tested on other animals. The remainder
of this paper reviews the history of this
line of research and some recent ad-
vances, suggests directions for future
research, and discusses the benefits of
chemical contraceptionin wild and feral
species.
HISTORY
Canidae
The use of antifertilitycompounds in
canids was promptedby the discovery in
1953that mismatedbitches could be in-
duced to resorb the embryos if treated
with the synthetic estrogen diethylstil-
bestrol (DES) (Jackson 1953). Linhart
(1963) proposed the use of steroid anti-
fertilityagents to control red fox (Vulpes
fulva) populationsand sylvatic rabies. A
year later, Linhart and Enders (1964)
demonstratedthatfemale foxes force-fed
50 mg of DES from nine days before
matingto ten days after mating became
infertile.DES apparentlyacted by caus-
ing implantationfailure or early embry-
onic death. The same dose of DES had
no significanteffects upon male foxes.
At about the same time, Balser (1964)
attempted to control coyote (Canis la-
trans)populationswith DES. Initiallab-
oratorytests had shown that DES given
just before or after mating inhibitedim-
plantation in coyotes just as it had in
foxes. Tallow baits filled with 100 mg of
DES were droppedin certain sections of
New Mexico. Female coyotes caught
several weeks later showed signs of re-
productive failure. Approximately 75%
of these females showed implantation
failureor fetal resorption.
There were, however, limitations in
the effectivenessof DES baitsfor inhibit-
ing fertilityin coyotes. First, DES inges-
tion had to be timed relatively precisely
in the coyote reproductivecycle for suc-
cess. This problemwas compoundedby
the fact that coyotes did not take nonliv-
ing baits with any regularityduring the
breedingseason. There also appearedto
be some problemin the absorptionof the
DES in tallow baits. Finally, Balser
(1964)reportedthat coyotes that did take
the DES-drugged bait appeared to re-
main sexually active longer than usual
486
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and were thoughtto be reproducinglater
in the season.
Since Balser's work with coyotes, re-
search on contraception in canids has
been redirected to domestic dogs. Tak-
ing advantageof advances in reproduc-
tive endocrinology, Simmons and
Hamner (1973) placed silicone rubber
implants containing testosterone or an-
drostenedionesubcutaneouslyin female
beagles. The bitches were kept in con-
stant anestrusfor 420 to 840 days; when
the implantswere removed, normalfer-
tility was restored. Estrus was clearly
suppressedin these dogs, but this study
did not identify the actual mechanismof
action. The most probable explanation
was that the androgenic steroids had
interfered with the hypothalamic-pitu-
A crucial issue in all
long-range wildlife
population management
programs is whether the
cure for the problem is
worse than the problem.
itaryaxis, blocked the release of luteiniz-
ing hormone (LH) and follicle stimulat-
ing hormone(FSH), and inhibitedfollicle
stimulation and/or ovulation. Unfortu-
nately, in addition to producinginfertil-
ity, the androgenicsteroids caused mas-
culinizationand altered the behavior of
the female in a mannerunacceptableto
dog owners.
Shortly thereafter,other investigators
explored using oral progestins for con-
trolling fertility in canids. Melengestrol
acetate, an oral progestin, was given to
bitches in doses of 200 J[g or more per
day, and all estrus activity was inhibited
for a treatmentperiod of 243 days (Soko-
lowski and VanRavenswaay 1976). The
positive results obtained with melenges-
trol acetate led to the first progestin
approved for commercial use in dogs.
The compound, megestrol acetate (Ova-
ban, ScheringCorp., Kenilworth,NJ), is
an effective oral contraceptive in the
bitch, highly reliable, and with few side
effects (Wildtand Seager 1977).At about
the same time in the Netherlands,anoth-
er syntheticprogestin, 14 alpha,17alpha-
propylidene-dioxy-progesterone,(proli-
gestone) was tested in dogs; it
suppressedestrus in 97% of the bitches
fed the compound (VanOs and Olden-
kamp 1978).
All use subject to JSTOR Terms and Conditions
Following these successes, attention
returned to androgenic steroids. The
synthetic androgenic steroid 17-beta-
hydroxy-7 alpha,17-dimethylester-4-en-
3-one (Mibolerone) given orally to
bitches for 240 days suppressed estrus
and probablyovulationfor the entire test
period without the objectionablemascu-
linization seen earlier with testosterone
(Sokolowski and Zimbelman1976).
Following the work of Linhart and
Enders (1964) and Balser (1964), Chea-
tum & Hansel1investigated the efficacy
of several different reproductive inhibi-
tors in a colony of red foxes duringfour
annual breeding seasons; three of the
compounds studied were found to be
effective. Various mixtures of clomi-
phene isomers [l-(p-beta-diethylamino-
ethoxyphenyl)-1,2, diphenyl-2-chloro-
ethylene] given at weekly intervals
throughoutthe breeding season did not
interferewith the occurrenceof estrus or
matingbut prevented pregnancies in all
vixens receiving it. The data suggested
that clomiphene may have impairedfer-
tilization, but the mechanism of action
was not described.
Diethylstilbestrolgiven in meat baits
on the day of mating or ten days there-
after prevented implantationin vixens.
However, if the DES was administered
in tallow, rather than meat, it lost its
efficacy. The choice of bait is clearly
important when attempting to deliver
synthetic steroids to carnivores. Chlor-
madinone acetate (6-chloro-6-17-ace-
toxyprogesterones) administered orally
every four, seven, or ten days prevented
estrus in most vixens until the feeding
regimenwas halted.
One of the most importantfeatures of
the Cheatum and Hansel study was an
attempt to inhibit male fertility. Sper-
matogenesis in male foxes was inhibited
by feeding them a mixture of DES and
chlormadinoneacetate in meat baits at
seven-day intervals for four weeks, be-
ginningthe firstweek in December. Five
weeks after drugadministrationwas dis-
continued, spermatozoawere present in
the seminiferoustubules but not in the
epididymides.No significantimpairment
of spermatogenesis was observed with
each of these compounds alone at the
same dosages.
The study of foxes by Cheatum and
Hansel and the study of Linhart and
Enders (1964) also demonstratedthat a
'E. L. Cheatumand W. Hansel. 1967. Rabies con-
trol by inhibitionof fox reproduction.Unpublished
ms., CornellUniversity, Ithaca, NY.
BioScience Vol. 35 No. 8
synthetic estrogen related to DES, mes- synthetic progestins could suppress es- in both cases there was significantfetal
tranol(3-methylether of ethinyl estradi- trus in female cats (Harrisand Wolchuk loss. The intramuscularapproach was
ol), was an effective reproductiveinhibi- 1963).The precise formof the compound more effective but clearly impracticalin
tor. No foxes fed MES for five days after and dose were refined, and by 1976 the animalsas secretive as white-taileddeer.
mating produced pups. Although most progestinmegestrol acetate was routine- A year later Bell and Peterle (1975)
first feedings of mestranol were readily ly administeredto domestic cats to sup- implanted both synthetic estrogen and
accepted, the vixens only nibbledat sub- press estrus (Burke 1977). Megestrolac- progestin in white-tailed does during
sequent drugged feedings. Thus it ap- etate was next applied to feral cat pregnancy. The implants were clearly
pears that MES could not be disguised populations in England with success superior to injected steroids; pregnan-
for prolonged administration to foxes (McDonald1980, Remfry 1978). cies ended in a significant number of
and probablynot to coyotes either. Seals et al. (1976) used injectableme- treateddoes. Matschke (1977a)adminis-
A somewhatdifferentapproachto ca- droxyprogesterone acetate and melen- tered DES orally to female white-tailed
nine fertility control was taken by Al- gestrolacetate in silastic implantsin cap- deer in an encapsulatedform. Doses of
Kafawi et al. (1974). This team attempt- tive African lions (Panthera leo), tigers 500-1000 mg encapsulatedin a modified
ed to immunize dogs against their own (Felis tigris), leopards (F. pardus), and gelatin interruptedpregnancy in a high
luteinizing hormone with injections of jaguars (F. onca). Long-term suppres- percentageof cases. However, poor ac-
human chorionic gonadotropin (hCG). sion of estrus resultedin complete inhibi- ceptance by the does, the need for very
The experiment failed because canine tion of reproduction,yet when the drugs high doses, and postabortion pregnan-
gonadotropinsand hCG did not cross- were removed, fertility was restored. cies led Matschke to conclude the ap-
react, but the concept was promising.A proach was not practical. Matschke
year later Faulkneret al. (1975)reported (1977b, c) then studied two synthetic
varying degrees of success in immuniz- In any wildlife fertility progestins, MGA (17 alpha-acetoxy-6-
ing dogs against LH through the use of methyl-16-methylene-4,6,pregnadiene-
antibodiesthat were considerably more program it is 3,20-dione) and DRC 6246 (17 alpha-
specific for canine LH. unreasonable and allyl-17-beta-hydroxy-3-oxoestra-4,9,1 1-
triene). Does were given 50 mg and 1.0 g
Aves probably unwise to daily doses of these drugs, respectively,
during the breeding season. Although
Antifertilityresearch in birds is some- attempt zero population does accepted both compounds without
what more limited. Davis (1959) demon- growth. difficulty, neither prevented or delayed
stratedthat testis weights and spermato- conception. Matschke (1980) hypothe-
genesis could be reduced in starlings sized that not enough of the steroids
(Sterna vulgaris) by feeding the com- Steroid fertility control in large cats is accumulatedin the animals' body fat to
pound triethylene melamine (TEM). He being appliedin Etosha National Parkin bring about prolongedaction.
then suggestedthat the broadconcept of Namibia,where Africanlion populations Roughton(1979), also realizingthe im-
populationcontrol throughchemical ga- are getting out of hand. In this case, practicalityof implantsin deer, fed 0.6-
metocidesbe appliedto wildlife manage- depot progestinsin silastic implantshave 1.0 mg of melengestrolacetate per head
ment (Davis 1961). Following up on the been placed in female lions in an effortto daily to white-tailed does during the
concept, Vandenberghand Davis (1962) slow reproduction(Chadwick 1983). breeding season. Reproduction was
showed that TEM could inhibit repro- Chanet al. (1981)took an immunologi- completely inhibited, there were no un-
duction in breeding populations of red- cal approachto fertility control in cats. toward side effects, and fertility was
winged blackbirds (Agelaius phoeni- They homogenized feline ovaries and later restored. To be effective, however,
ceus). Testis weights were reduced in raised rabbit antibodies against them. the drughad to be given daily, a require-
TEM-treated males, and meiosis was The antibodies, when administered to ment that may be difficultto meet in wild
also inhibited; however, the drug's ex- pregnantcats, caused some fetal resorp- populations.
treme toxicity precludedpracticalappli- tion, but results were in general discour-
cation. Followingthis, Elder(1964)dem- aging. Once again, as in dogs, nonspeci-
onstrated that Provera, an oral ficity in the antibodyappearedto be the Rodentia
progestin;Arasan, a common fungicide problem. The concept of chemical fertility con-
and seed disinfectant; and the anticho- trol for managing pest rodent popula-
lesterol agent SC-12937(22,25-diazacho- tions was originallyput forth by Howard
lestanol dihydrochloride) all brought Cervidae (1967). Marsh and Howard (1969) fed
about reversible inhibition of ovulation There has been some interest in fertil- 0.05%mestranolbaits to wild rats (Rat-
in pigeons (Columba livia). More recent- ity control in ungulates also. Between tus norvegicus) and observed an immedi-
ly, Lacombeet al. (1984)showed that the Decemberand March,Greeret al. (1968) ate and significantreductionin pregnan-
compound ornitrol (20,25-diazacholes- administeredDES intramuscularlyto 36 cies. Poor bait acceptance of the
terol dihydrochloride) inhibited testes pregnant cow elk (Cervus canadensis) synthetic estrogen gave short-lived re-
growth in red-winged blackbirds, but from the northernYellowstone National sults. Following this study, Howard and
only if it was fed at precise times in the Parkherd. Doses of 75-200 mg terminat- Marsh (1969) and Storm and Sanderson
reproductivecycle of the birds. ed pregnancyin 30% of the treated ani- (1970) expanded this line of research to
mals. Following this, Harder (1971) and include voles (Microtus californicus, M.
Felidae Harder and Peterle (1974) fed DES to montanus, and M. pennsylvanicus).
female white-taileddeer (Odocoileusvir- Again, rats did not accept mestranol
It had been shown as early as 1963that ginianus) before and duringpregnancy; baits well, and doses had to be reduced

September 1985 487

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 to 0.005% before voles would accept tissue encapsulation and interference
species where harem-like groups exist,
baits regularly.Reproductionwas inhib- with drug release.
greaterefficiencycould come from caus-
ited in the voles, and pups receiving Another exciting advance, which we
ing infertilityin the male, provided that
mestranol through the mothers' milk used in ourferalhorse program(Kirkpat-
sexual behavior were not significantly
never developed sexually, becoming ir- altered. rick et al. 1982, Turnerand Kirkpatrick
reversiblysterile. Recently Kirkpatrick et al. (1982), 1982),is the developmentof a biodegrad-
Brooks and Bowerman (1971) and Turner and Kirkpatrick (1982), and able encapsulation process to permit
Mischler et al. (1971) evaluated a new Turner(1984), reported successfully in-
long-termsustained release of injectable
powerful estrogenic compound, 17 al- hibiting reproduction in feral horsescontraceptiveagents (Beck et al. 1980).
pha-ethynylestradiol-3-cyclopentyl (Equus caballus) by lowering sperm One successful form of microencapsula-
ether, later known as Quinestrol. This counts in stallions. A microencapsulated tion, the DL-lactide compound, permits
compoundwas shown to be more effec- form of testosterone propionatewas in- single injections to sustain a release of
tive than mestranolfor antiovariantreat- jected into stallions several months be- contraceptive steroids for up to several
ment in rodents. Doses as low as 10 Ixg fore the April-June breeding season in years. Such a system would be invalu-
completely inhibited reproduction, but Challis, Idaho. The poly (DL-lactide) able in wildlife management.
once again poor bait acceptance in rats coating permitted a sustained release, Recent advances in our knowledge of
made the approachimpractical. thereby causing oligospermia and im- male fertility control, coupled with new
At present, only one drug has been pairmentof sperm motility for up to six antifertilityagent delivery systems, pre-
successfullytested as an antifertilityagent
months. The stallions' behavior was un- sent a whole new dimension to fertility
in rodents and carries FDA approvalfor control in wild species. We have suc-
that use. Ericsson (1975) fed Norway cessfully used testosterone alone to sup-
rats alpha-chlorohydrin (3-chloro-1,2- sperm production in feral horses
propanediol).The drugwas lethal in both
Two of the more obvious press
(Kirkpatrick et al. 1982, Turner and
sexes if ingested in high enough quanti- research needs are Kirkpatrick1982); combinations of ste-
ties (LD50 164 mg/kg body weight) but roids may be even more effective. Dana-
also caused permanentsterility in male bait acceptance by small zol, the 2,3-d isoxazol derivative of 17-
rats at sublethaldoses. The drug causes mammals and using ethinyl testosterone, is very effective in
a blockageat the head of the epididymis, reducingsperm counts in men, and pro-
which prevents spermatozoafrom leav- long-acting injectable gestin-testosterone enanthate combina-
ing the testis. The sterilizedmale rats are steroids in large tions now under study also appeareffec-
otherwise healthy and mate normally, tive in reducing sperm numbers to the
althoughwithout results. mammals. level of infertility(Steinberger1980).All
Once alpha-chlorohydrinenters the these compounds have reversible anti-
rat, normalmetabolism,bacterialdegra- fertilityeffects, can be coupled with new
dation, and contact with water reduces affected and breeding took place, but delivery systems such as biodegradable
the drug to water, CO2, and chloride; there was an 83%reduction in foal pro- capsules, can be deliveredremotely, and
nontarget species that might catch and duction. In addition, Kirkpatricket al. appearto have no serious side effects.
eat a drugged rat will thus suffer no (1982)demonstratedthat repeated injec- So far we have confined our discus-
consequences. The drug, now marketed tions of testosterone cypionate and sin- sion of contraceptive agents almost ex-
commercially (Epibloc, Pestcon Sys- gle injections of Quinestrol could also clusively to steroids. The Nobel Prize-
tems, Inc., Alhambra, CA) is microen- cause oligospermia in stallions; these winning work of Andrew Schally and
capsulated in a vinyl resin-based wall two drugshave not yet been field tested, Ralph Guillemin on peptide releasing
material, and bait acceptance has not however. hormones in the hypothalamus opens
been a problem. Because it is toxic and entirely new doors to fertility control.
its antifertilityeffects are permanent,al- Synthetic analogs of luteinizing hor-
RECENTADVANCES
pha-chlorohydrinhas limiteduse in wild- mone-releasinghormone (LH-RH) have
life managementbut is clearly useful for Froma scientificviewpoint, the future been administeredby injection and have
controllingcertain pest rodent species. for fertility control in wild animalpopu- been extremely effective in blocking
Recently, Garrettand Franklin(1983) lations is bright. Long-acting injectable ovulation (Schally 1983).
successfully inhibited reproduction in contraceptivesofferthe wildlife manager Recently Vickery et al. (1984)success-
black-tailedprairiedogs (Cynomysludo- some of the strongestpopulationcontrol fully suppressed fertility in male dogs
vicanus)in Wind Cave National Parkby measures, largely because these drugs with repeated injections of an LH-RH
feeding DES-treated oats (0.11% active can be administeredremotely with tran- agonist. The agonist, [D-Nal(2)6-LH-
ingredient).Reproductiveinhibitionwas quilizer guns, thereby avoiding expen- RH] was administeredintramuscularlyin
almost 100% effective, and there was sive capture and handling programs. doses of 10 pLg/kg,and testis volume,
total reversibility when drug treatment Some 25-30 differentcommercialinject- ejaculate volume, sperm count, and
was withdrawn. able contraceptive steroids are now spermmotilityall declined for periods as
available, and several new long-acting long as 172 days. Libido was depressed
progestins show greatpromise. Although but could be restored by testosterone
Equidae
subdermal steroid implants have been implants. The results of this work were
With a few notable exceptions, the tested with some success, the newer encouraging,and furtherresearch is un-
concept of managingwild andferalpopu- injectablelong-acting steroids appear to der way.
lations through fertility control has fo- work better; the steroid subdermalim- One of the most exciting possibilities
cused on females. However, in certain plants quite often result in connective for fertility control lies in immunology.

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Raising antibodies against sperm (Tung fertility is an easy proposition if resist- worse than the problem. The biosystem
1976)and the zona pellucida of the em- ance to bait acceptance can be over- within which each target species dwells
bryo (Glass and Hanson 1974)has been come. Encapsulating steroids in a dis- is dynamic and flexible within certain
possible for some time. Since gonado- guised and acceptable medium may limits. Unfortunately, in most cases we
tropic hormones, such as follicle stimu- represent a profitabledirection for new have not yet delineated those limits and
lating hormone (FSH) and luteinizing research. For example, Calanchi (1976) must work on the principle of minimal
hormone (LH), and the hypothalamic made 5-mg doses of steroid palatableto perturbance.This means extensive pre-
releasinghormones,such as LH-RH, are human patients by microencapsulating liminary studies to establish normal re-
all peptides, the molecules lend them- them; the drugs had been unacceptable productive and behavioral patterns and
selves to possible immunologicalattack. in uncoated 200-jig doses. then furtherstudies to assess treatment
Immunizationto monkey FSH (Niesch- Although we successfully delivered effects. Such studies must encompass
lag and Wickings 1981), rodent LH-RH long-acting contraceptive steroids by complete pharmacologyof the drugs be-
(Fraser 1975), and canine LH (Faulkner darts from a tranquilizergun (Kirkpat- ing used, including specific side effects,
et al. 1975)has been accomplishedwith rick et al. 1982, Turnerand Kirkpatrick catabolism,andpossible biologicalactiv-
varyingdegrees of success. Practicalap- 1982),the technology we used was rela- ity of the excreted metabolites that may
plications of this approach include im- tively crude. All the newer injectable be ingested by other species in the
munizing sheep against LH and cattle progestin and androgensteroids require habitat.
against LH-RH (Robertsonet al. 1982). testing in a wide arrayof large wild and Also crucial is the extent to which
Currently,however, the technology in feral species and in both male and female drugsinterferewith libido and sociosex-
immunologicalfertility control is not at ual behavior.In any wildlifefertilitypro-
the point where widespread application gram it is unreasonable and probably
to wild species is possible. One problem unwise to attempt zero population
is that immunizationto LH or LH-RH in Wildlife-its preservation growth.If males are the targetand only a
males leads to a loss of libido as well as and particularly its percentage will be made infertile, then
suppressionof spermatogenesis. Immu- the antifertilityagent must not interfere
nizationagainstFSH may have potential management-is more with libido or other behavior that con-
value in males, without significant often than not a highly tributes to their success in mating. An-
other important consideration is the
changesin sexual behavior, and immuni-
zation against all three peptides men- charged emotional issue. long-termeffects of behavioral changes
tioned above may be effective in the on social organization. Any potential
female. This form of contraceptioncan antifertilityagent must be screened for
last for as long as one to three years animals. In addition, biodegradablemi- direct and indirect behavioral effects.
(Nieschlag and Wickings 1981) and in croencapsulationshould be added to the The choice of behavioralendpoint(s)for
most cases is reversible. One of the growinglist of injectablesteroids to pro- such screening is critical; sexual behav-
major hurdles to effective use of the vide a larger spectrum of long-acting ior and at least one other behavior in-
immunological approach has been the contraceptives. volved in maintaining social structure
failure to obtain sufficient quantities of If chemical fertility control is to be would be desirable.
pureantibodiesand species-specificpep- successful, the effects of contraceptive The lessons learnedover the past dec-
tide hormones (Wildt and Seager 1977). steroids on the length of the target spe- ades of research on controlling human
The current advances in monoclonal cies' breeding season must be carefully fertility must not be lost to those who
antibody production by hybridomas determined. Polyestrus seasonal breed- wish to apply this technology to wild
(Goding 1980) may help solve this ers might well remain infertile through- species. Regardlessof the safety or effi-
problem. out a spring and early summerbreeding cacy of a chemicalfertilitycontrol agent,
Still another potentially useful ap- season, but care must be taken that the there are importantpublic issues to be
proachis using plant productsthat inter- reproductiveprocess is not just delayed. considered and political barriers that
fere with reproduction.A recent review A populationwhose reproductiveactivi- must be overcome before widespread
(Farnsworthand Waller 1982) listed 50 ty normallyconcludes in summerand is success can be achieved. Wildlife-its
plant families, genera, and species that delayed to the fall will produce young preservationandparticularlyits manage-
have documented antifertilityeffects in thatwill probablynot survive the winter. ment-is more often than not a highly
males and females. Using some of these Such a situationwould be inhumaneand charged emotional issue. Thus, a final
plants, particularlyin managing repro- unacceptable. thrust for future research should be
duction in herbivores, deserves further The technology for practicalimmuno- aimed at determiningwhether the public
consideration. logicalcontrollags far behindthe contra- will accept fertility control in wild spe-
ceptive steroid approach, but interrupt- cies and how to change resistance to
ing reproductionwith antibodies offers acceptance.
DIRECTIONS FOR FUTURE greater species-specific mechanisms of
action and virtuallyno risk to nontarget BENEFITS
RESEARCH
species. The production of pure mono-
Two of the more obvious research clonal antibodies by means of hybrid- Perhaps the most compelling reasons
needs are bait acceptanceby small mam- omas would represent an important for using chemical fertility control are
malsand usinglong-actinginjectableste- advancement. social. Simply, the approachis humane,
roidsin largemammals.Researchresults A crucial issue in all long-rangewild- andpublicacceptanceis morelikely than
to date make it clear that the control of life populationmanagementprogramsis in the case of hunting, poisoning, or
rodent, canine, and other small animal whether the cure for the problem is trapping. Not only is individual animal

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discomfort minimized or eliminated, but
there is an often-overlooked secondary
humane aspect. Where hunting is forbid-
den by law and relocating animals is
economically or physically unfeasible,
overpopulation is likely to end in disease
and death by starvation. Fertility control
is more likely to be permitted within
such protected areas.
Depending on the species involved,
chemical fertility control can also bring
economic advantages. Consider, for ex-
ample, the cost of removing a feral horse
from public lands in the United States:
$600 to $1000 per animal (Rey 1975). Not
only is this initial cost substantial, but
the population will increase every year,
necessitating an annual reduction pro-
gram. The initial cost of chemical fertil-
ity control is comparatively low per ani-
mal treated, and in cases where the male
of a polygamous species is the target, the
cost is reduced in a way proportionate to
the degree of the polygamy. In feral
horses, a single treated stallion may ulti-
mately inhibit reproduction in three to
ten mares. There are long-term econom-
ic benefits as well, since well-planned
fertility control programs will prevent
unnecessarily large population in-
creases, thereby reducing the magnitude
of future control programs.
Chemical fertility control is a flexible
management tool, permitting a large va-
riety of control manipulations. With
smaller mammals like skunks (Mephitis
mephitis) and drugged baits, local popu-
lations can be controlled with some pre-
cision, rather than subjecting entire
states or districts to a single approach.
Where drugs can be delivered remotely
to large animals, particular herds, bands,
and even individuals can be singled out.
The reversibility of the drug action adds
even more management flexibility. A
large and unanticipated winter kill, for
example, can be offset by withdrawing
drug treatment for one or more breeding
seasons. Since breeding animals have
not been permanently removed, herd or
population size can be restored to safe
levels.
While the social advantages of chemi-
cal fertility control are impressive, the
biological ones are exceptional. Remov-
ing an animal from a population by hunt-
ing, trapping, poison, or relocation is
permanent; the genes are lost from the
pool forever. Because chemical contra-
ception is reversible, its use within an
intelligent management plan keeps the
gene pool intact. This may be an ex-
tremely valuable concept where dwin-
dling habitat results in localized over-
490
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population of rare or endangered
species.
Another obvious advantage is the abil-
ity to concentrate upon target species
without serious damage to other animals.
Delivering bait to a particular species
without having a variety of other animals
ingesting the drug is probably impossi-
ble, but through careful manipulation of
drug types and dosages, effects on non-
target species can be minimized. A dose
of a particular contraceptive steroid that
inhibits vole reproduction may well have
no effect on foxes or eagles; more re-
search is needed before this can be stat-
ed with certainty.
Moreover, the peak breeding season
for one species often varies significantly
from that of another, which makes it
easier to deliver bait without interfering
with nontarget species. Red foxes, for
example, breed from December to
March, but skunks do not begin breeding
until March (Asdell 1964). Thus, a bait-
delivered fertility control program could
be tailored for skunks to avoid interfer-
ence with fox reproduction. At very
least, accidental ingestion will not kill
nontarget species or, except for extreme-
ly young animals, cause irrevocable fer-
tility loss.
Finally, chemical fertility control af-
fords an approach that, when properly
evaluated, will not influence the social
structure of the animal populations in-
volved. Since sexually mature individ-
uals are not actually removed, or for that
matter even harrassed, the hierarchy of
the population is not altered unwittingly
by human intervention.
Controlling overabundant wildlife
populations through contraception is a
potentially powerful management tool
that has received surprisingly little atten-
tion. Continued human encroachment on
critical habitat, coupled with increased
public resistance to traditional control
programs, will ultimately require new
solutions to overpopulation problems.
Much of the scientific knowledge neces-
sary to provide safe, effective, and hu-
mane control is already at hand; wildlife
managers must be bold enough to seek
these new directions.
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1976.Evaluationof selected compoundsfor earth.
estrus controlin the bitch. Am. J. Vet. Res. The ocean'spoundingsurf,Spring
37: 939-941.
coveringthe earthwith that season's
Steinberger, E. 1980. Current status of re- newness,a bird'ssong reaching
search on hormonal contraception in the out... and we are moved.
male. Res. Front. Fertil. Regul. 1: 1-12. Our curiositycalls us. We explorea bit.
Storm, G. L., and G. C. Sanderson. 1970. Some travelto a distantwilderness,some
Effectof mestranolanddiethylstilbestrolon no fartherthan a comer of the back yard.
captive voles. J. Wildl. Manage. 34: 835- But each of us looks and listensto the
843. wonderof the naturalworldaroundus.
Tung, K. 1976.Antifertilityvaccines: consid- We enjoythe earth'sbeauty,it's
erationsof theirpotentialimmunopatholog- grandeur.The endless sweep of colors,
ic complications.Int. J. Fertil. 21: 197-206. and sounds, and everywhere,the
Turner, J. W., Jr. 1984. Given a free rein excitementof life.
prolific mustangs gallop into trouble. But there are some that do not see the
Smithsonian14(11):88-96. wonderof it all. And so we, of the Sierra
Turner, J. W., Jr., and J. F. Kirkpatrick. Club,join togetherto protectthe earth.
1982. Androgens, behavior and fertility We workfor legislationthat guarantees
control in feral stallions. J. Reprod. Fertil. clean air and water.To regulatethe use
Suppl. 32: 79-87. and disposalof poisonoustoxic chemicals.
Vandenbergh,J. G., and D. E. Davis. 1962. To set aside the most specialplacesfor
Gametocidaleffects of triethylenemelamine parksand wilderness.
on a breeding population of red-winged We inviteyou to join with us.
blackbirds.J. Wildl.Manage. 26: 366-371. To explore,to enjoy.To protectthis
VanOs, J. L., and E. P. Oldenkamp. 1978. wondrousearth. For all of us... forever.
Oestrus control in bitches with proliges- For membershipinformation,write
trone, a new progestational steroid. J. SierraClub,530 Bush Street, San
Small Anim. Pract. 19: 521-529. Francisco,CA 94108, (415) 981-8634.
Vickery,B. H., G. I. McRae, W. Briones, A.
Worden,R. Seidenberg,B. D. Shanbacher,
and R. Falvo. 1984. Effects of an LHRH Sierra
agonistanalogupon sexual functionin male
dogs. J. Androl. 5: 28-42. Club
Wildt, D. E., and S. W. J. Seager. 1977. Re-
productioncontrol in dogs. Vet. Clin. N.
Am. 7: 775-787.
491
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