Zoo Biology 17:369371 (1998)

EDITORIAL

Delayed Reproduction:
The Good and the Bad
There is nothing like a failed reproductive effort with high-profile exotics to
focus ones attention on the search for possible contributing factors. We refer, in this
instance, to San Diegos pair of giant pandas, from which staff has tried valiantly to
coax mating in the past two seasons. Imported in 1996 from a captive facility in
China for scientific study, the spontaneous occurrence of a strong female estrus in
each of the two past mating seasons provided an opportunity to document how male
and female communicate during one of the major instances of social contact in this
predominantly solitary mammal.
The male of this pair was confirmed to be about 1920 years of age during the
first seasonan age beyond which very few pandas in the wild survive, according to
experts such as George Schaller and Pan Wenshi. Our data reveal that failure to mate
in both seasons was clearly due to a motivational deficit in this aged individual. To
state it plainly, the old fellow was steadfastly unmoved by the avid solicitations of
his young companion.
On testing, we found this males semen parameters to be excellent and his
testosterone levels normal. He is in reasonably good health, certainly vigorous enough
to perform sexually. He lives apart from the female, as do pandas in the wild, but she
is not a total stranger to him. His olfactory sense appears to be intact, and he was
given ample opportunity to investigate her scent and urine marks as she approached
estrus. Having lived his first decade or so in the wild, one would expect this male to
know what to do in the presence of a highly solicitous female. Too old to become
aroused? The literature on sexual senescence in mammals would certainly indicate
that this is a real possibility.
A different kind of scenario is suggested by the case of female cheetahs in our
collection. A healthy youngster named Jillani began to show signs of estrus shortly after
pubescence, mated, conceived, and gave birth to the first of several litters. As fate would
have it, in her prime years this female was caught up in a national health moratorium and
was precluded from further breeding. She remained in good health and continued to
cycle to the end of her life. Tamu, another female in the collection, also became interested in males shortly after pubescence, but from this time was under a different kind of
moratorium, that of the genetically surplus animal. Gradually, her cycles waned and eventually ceased altogether. Today, as a candidate for breeding, she is unresponsive, and
hormonal analysis shows that her ovaries have become quiescent.
1998 Wiley-Liss, Inc.

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Editorial

Obviously, a couple of anecdotes do not make a definitive case. However, we

have been involved with enough species in zoos to know that delayed reproduction
is a fairly common occurrence and that a great deal of effort is being directed at
trying to get offspring from genetically valuable but reproductively aged individuals.
Virtually every master plan has a number of individuals that are on the borderline of genetic need or at least are not needed for the moment, so are held in reserve
for a later time when breeding opportunities might at last arise. Others are proposed
for more widely spaced breeding, that is, at intervals greater than the average interbirth
interval, as a way of extending generation times and thereby reducing the rate at
which genetic diversity is lost. Still others belong to a class that, for a variety of
reasons, has reached their later years without being afforded an opportunity to reproduce. Anyone who has been involved in master planning even tangentially is aware
of the measures that are advocated in the interest of conserving genetic diversity in
zoos small, closed populations. Although important genetic objectives are thereby
served, managers operating in the real world often find that it is not as easy as one
might think to get the reproductive juices flowing again. Whether it is strictly an age
effect or some combination of age and withheld opportunities for breeding that exacerbates the problem remains to be determined. Yet, there are abundant data to indicate that relegating reproduction to the later stages of reproductive life entails a cost,
particularly in the case of females.
We can easily agree that the ravages of passing time will have their effect on structures and systems, leading to declining fertility, declining interest, and declining performance. The supply of germ cells diminishes or may become exhausted, hormonal balances
change, tissues become flaccid, disease turns the normal into the pathological, and the
waking hours become more focused on basic needs for coping and survival than on
procreative activity. Age effects, however, may not be the whole story.
Why do the ovaries from aged multiparous mice yield fewer histological abnormalities than those of same-aged virgins? How is it that a lifetime of barrenness
results in lower rates of embryo recovery and an increased percentage of abnormal
embryos in maiden versus foaling mares? Why should the incidence of polyps, endometrial pathology, positive uterine cultures, and generally poor uterine health be
higher in maidens than in same-aged dams? Apparently, quite apart from the increased risk of debilitating infections as the individual ages, there is a price to pay
for a lifetime of reproductive inactivity as well.
Horse breeders have long lamented the negative effects of a phenomenon known
as the maiden mare syndrome on breeding. In simple terms, it is often more difficult to obtain breeding in later life from mares who have never bred than from likeaged mares that have been reproductively active. In all probability, if we took the
time to look, we would find the maiden syndrome extends to more than just horses,
perchance even to those non-domesticated forms that we desperately want to conserveelephants, rhinos, gorillasand many more. It seems possible that we as
master planners, in advocating that reproduction be delayed to the later years, may
be imposing on managers, clinicians, and researchers a heavier burden than we know.
Ours is a profession in which we embrace the human can do spirit, as seen in
efforts intended to bring wild forms back from the brink of extinction. We participate as specialists in various disciplines in a spirit of collegiality and hopefulness but
experience despair when these almost super-human efforts fail. Our sense is that the
geneticists and the demographers, in developing models intended to guide reproduc-

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tive efforts in essential ways, have probably gotten a large part of the drama correct.
Increasingly, however, we are learning that strategies implemented without due allowances for the managers realm of operation entail costs, often hidden and rarely
documented, that predispose the effort to fail. It is the living, breathing organism that
must propagate itself, and disciplines concerned with this side of the equation have a
lot of catching up to do if self-sustaining populations are to become a reality.
Donald G. Lindburg, Editor
Barbara S. Durrant
Zoological Society of San Diego