Much of what is known of the length of life of animals other than man derives from observations of domesticated species in laboratories and zoos. One has only to consider how few animals reveal their age to appreciate the difficulties involved in answering the apparently simple question of how long they live in nature. In many fishes, a few kinds of clams, and an occasional species of other groups, growth is seasonal, so that annual zones of growth, much like tree rings, are produced in some part of the organism. Among game species, methods of determining relative age by indicators such as the amount of tooth wear or changes in bone structure have yielded valuable information. Bird bands and other identifying marks also make age estimation possible. But one of the consequences of the fact that animals move is that very little is known about the life span of most species as they exist in nature.
Maximum and average longevity
Many of the extreme claims of longevity that are occasionally made for one species or another have consistently been proven false when subjected to critical scrutiny. Although the maximum life span that has been observed for a particular species cannot be considered absolute, since a limited number of individuals at best has been studied, this datum probably provides a fair approximation of the greatest age attainable for this kind of animal under favourable conditions. Animals in captivity, which provide most of the records of extreme age, are exposed to far fewer hazards than those in the wild.
Environmental influences
Life span usually is measured in units of time. Although this may seem eminently logical, certain difficulties may arise. In cold-blooded animals in general, the rate of metabolism that determines the various life processes varies with the temperatures to which they are exposed. If aging depends on the expenditure of a fixed amount of vital energy, an idea first proposed in 1908, life span will vary tremendously depending on temperature or other external variables that influence life span. There is considerable evidence attesting at least to the partial cogency of this argument. So long as a certain range is not exceeded, cold-blooded invertebrates do live longer at low than at high temperatures. One of the most striking examples of this phenomenon occurs in Greenland sharks (Somniosus microcephalus), where some individuals reach ages of 500 years or more. Rats in the laboratory live longest on a somewhat restricted diet that does not permit maximum metabolic rate. Of perhaps even greater significance is the fact that many animals undergo dormant periods. Many small mammals hibernate; a number of arthropods have life cycles that include periods during which development is arrested. Under both conditions the metabolic rate becomes very low. It is questionable whether such periods should be included in computing the life span of a particular organism. Comparisons between species, some of which have such inactive periods while others do not, are dangerous. It is possible that life span could be measured more adequately by total metabolism; however, the data that are necessary for this purpose are almost entirely lacking.
Length of life is controlled by a multitude of factors, which collectively may be termed environment, operating on a genetic system that determines how the individual will respond. It is impossible to list all the environmental factors that may lead to death. For analytical purposes it is, however, useful to make certain formal separations. Every animal is exposed to (1) a pattern of numerous events, each with a certain probability of killing the individual at any moment and, in the aggregate, causing a total probability of death or survival; (2) climatic and other changes in the habitat, modifying the frequency with which the various potentially fatal events occur; and (3) progressive systemic change, inasmuch as growth, reproduction, development, and senescence are characteristics intrinsic in the organism and capable of modifying the effects of various environmental factors.
Patterns of survival
Consider a group of similar animals of the same age. Although no two individuals can have precisely the same environment, let it be assumed that the environment of the group remains effectively constant. If the animals undergo no progressive physiological changes, the factors causing death will produce a death rate that will remain constant in time. Under these conditions, it will take the same amount of time for the population to become reduced to one-half its former number, no matter how many animals remain at the beginning of the period considered. The animals therefore survive according to the pattern of an accident curve. This is the sense in which many of the lower animals are immortal. Although they die, they do not age; how long they have already lived has no influence on their further life expectation.
Another group of animals may consist of individuals that differ markedly in their responses to the constant environment. They may be genetically different, or their previous development may have caused variations to arise. Those individuals that are most poorly suited to the new environment will die, leaving survivors that are better adapted. The same result can also be achieved in other ways. If the environment varies geographically, those individuals that happen to find areas in which existence can be maintained will survive, while the remainder will die. Or, as a result of their own properties, animals in a constant environment may acclimate in a variety of ways, thus adjusting to the existing conditions. The pattern of survival that results in each of these cases is one in which the death rate declines with time, as illustrated by the selection–acclimation curve.
In the absence of death from other causes, all members of a population may exist in their environment until the onset of senescence, which will cause a decline in the ability of individuals to survive. In a sense they can be considered to wear out as does a machine. Their survival is best described by individual differences among members of the population that determine the curvature of the survival line (wearing-out curve). The more the population varies, the less abrupt is the transition from total survival to total death.
Under the actual conditions of existence of animals the three types of survival (accident pattern, selection–acclimation pattern, wearing-out pattern) above all enter as components of the realized survival pattern. Thus in animals that are carefully maintained in the laboratory, survival is approximately that of the wearing-out pattern. Environmental accidents can be kept to a minimum under these conditions, and survival is almost complete during the major part of the life span. In all known cases, however, the early stages of the life span are characterized by a noticeable contribution of the selection–acclimation pattern. This must be interpreted as a result of developmental changes that accompany the early life of the individuals and of selective processes that operate on those organisms whose genetic constitutions are ill fitted for that environment.
In some of the larger mammals in nature, the existing evidence points to a similar survival pattern. In a variety of other animals, however, and including fishes and invertebrates, mortality in the young stages is so high that the selection–acclimation curve predominates. One estimate places the mortality of the Atlantic mackerel during its first 90 days of life as high as 99.9996 percent. Since some mackerel do live for several years, a mortality rate that decreases with age is indicated. Similar considerations probably apply to all those animals that have larval stages that serve as dispersal mechanisms.
When the postjuvenile portion of the life span is considered by itself, a number of animals for which such information has been gathered—including primarily fishes and birds—have survivorship curves that are dominated by the accident pattern. In these species in nature, death from old age apparently is rare. Their chance of surviving to an advanced age is so small that it may be statistically negligible. In modern times, human predation is a large factor in the mortality of these species in many cases. Since deaths from fishing and hunting are largely independent of age, once an animal has reached a certain minimum size, such a factor only makes the survival curve steeper but does not change its shape. One consequence of such increased mortality is that fewer old and large individuals are noticed in a population.
More complex survival patterns, such as the hypothetical one illustrated, undoubtedly exist. They should be looked for in those species in which extensive reorganization of the animal is part of the normal life cycle. In effect, these animals change their environment radically, in some cases several times during a lifetime. The frog offers a familiar example. During its period of early development and until shortly after hatching, the animal is subject to major internal, and some external, change. As a tadpole it is adjusted to an aquatic, herbivorous life. The metamorphosis to the terrestrial, carnivorous adult form is accompanied by varied physiological stresses that must be expected to produce a temporary increase in mortality rate. In some insects the eggs, larvae, pupae, and adults are exposed to and respond to quite different environments, and a survivorship pattern even more complex than that described by the composite curve may exist.
The same species will exhibit changed survival in different environments. In captivity an animal population may approach the wearing-out pattern; in its natural habitat survivorship may vary with age in a quite different way. Although one can assign a maximum potential life span to an individual—while realizing that this maximum may not be attained—it is impossible to specify the survivorship pattern unless the environment is also specified. This is another way of saying that life span is the joint property of the animal and the environment in which it lives.