Читать книгу The Prolongation of Life: Optimistic Studies - Élie Metchnikoff - Страница 10
I
THEORIES OF LONGEVITY
ОглавлениеRelation between longevity and size—Longevity and the period of growth—Longevity and the doubling in weight after birth—Longevity and rate of reproduction—Probable relation between longevity and the nature of the food
The duration of the life of animals varies within very wide limits. Some, as for instance, the males of certain wheel animalculæ (Rotifera) complete their cycle of life from birth to death in 50 or 60 hours, whilst others, like some reptiles, live more than 100 years, and quite possibly may live for two or three centuries.
Enquiry has been made for many years as to whether there are laws governing these different durations of life. Even the most casual observation of domesticated animals has shown that, as a general rule, small animals do not live so long as large ones; mice, guinea pigs, and rabbits for instance, have shorter lives than geese, ducks, and sheep, whilst these again are survived by horses, deer, and camels. Of all the mammals which have lived under the protection of man, the elephant is at once the largest, and the most long-lived.
However, it is not difficult to show that there is no absolute relation between size and longevity, since parrots, ravens, and geese live much longer than many mammals, and than some much larger birds.
As a general rule it may be said that a large animal takes more time than a small one to reach maturity, and it has been inferred from this that the length of the periods of gestation and of growth were in proportion to the longevity. Buffon28 long ago stated his opinion that the “total duration of life bore some definite relation to the length of the period of growth.” Therefore, as the period of growth is, so to say, inherent in the species, longevity would have to be regarded as a very stable phenomenon. Just as any species has acquired a fixed and practically invariable size, so it would have acquired a definite longevity. Buffon, therefore, thought that the duration of life did not depend on habits or mode of life, or on the nature of food, that, in fact, nothing could change its rigid laws, except an excess of nourishment.
Taking as his standard the total period of development of the body, Buffon came to the conclusion that the duration of life is six or seven times that of the period of growth. Man, for instance, he said, who takes 14 years to grow, can live 6 or 7 times that period, that is to say, 90 or 100 years. The horse, which reaches its full size in 4 years, can live 6 or 7 times that length of time, that is to say from 25 to 30 years. The stag takes 5 or 6 years to grow, and reckoned in the same way, its longevity should be 35 to 40 years.
Flourens29 although supporting his principle, thought that Buffon had been inexact in calculating the period of growth. In his opinion a better result can be obtained by taking the limit of growth as that age at which the epiphyses of the long bones unite with the bones themselves. Using such a mode of computation, Flourens laid down that an animal lived 5 times the length of its period of growth. Man, for instance, takes 20 years to grow, and he can live for 5 times that space, that is to say, 100 years; the camel takes 8 to grow, and lives 5 times as long, i.e., 40 years; the horse, 5 to grow, and lives 25 years.
However, even if we consider only the mammalia, it is impossible to accept Flourens’ law, without considerable reserve. Weismann30 has referred to the case of the horse, which is completely adult at 4, but lives not merely 5 times that period, but 10 or even 12 times. Mice grow extremely quickly, so that they are able to reproduce at the age of 4 months. Even if we take 6 months as their period of growth, their longevity of 5 years is twice as long as it would be according to the rule of Flourens. Amongst domesticated animals, the sheep is slow in reaching maturity; it does not acquire its adult set of teeth until it is 5 years old, and cannot be regarded as adult until then. None the less, at the age of 8 or 10 years, it loses its teeth and begins to grow old, whilst by 14 it is quite senile.31 The longevity of the sheep, therefore, is not quite three times its period of growth.
If we turn to other vertebrates, the variations in the relation of growth and the duration of life are still greater. Parrots, for instance, the longevity of which is extremely great, grow very quickly. At the age of 2 years, they have acquired the adult plumage and are able to reproduce, whilst the smaller species are in the same condition at the age of one. Incubation, moreover, is very short, not more than 25 days, and in some species not three weeks. None the less, parrots are birds which enjoy a quite remarkable longevity. The incubation period of domestic geese is 30 days, and their period of growth is also short. However, they may reach a great age, cases of 80 years and of 100 years being on record. In contrast with these, ostriches, the incubation period of which is 42 to 49 days, and which take 3 years to become adult, have a relatively short life.
H. Milne-Edwards32 many years ago contended that there was no importance in the supposed law of relation between gestation and longevity. He sums up his criticism as follows: “Although the period of uterine life is longer in the horse, that animal does not live so long as a human being; and some birds, the incubation of which only lasts a few weeks, can live more than a century.”
Bunge33 has recently taken up the study of the relations between the duration of growth and longevity, and has suggested a new means of investigation. He has observed that the period in which the new-born mammal doubles its weight is a good index of the rapidity of its growth. He has shown that whilst a human child requires 180 days to reach double its weight at birth, the horse, the longevity of which is very much less, doubles its weight in 60 days; a calf takes only 47 days for this; a kid 15 days; a pig 14 days; a cat 9–½; and a dog only 9 days. Although these facts are very interesting, the exceptions are too great to make it possible to base a law of longevity upon them. The period of weight-doubling in the horse is nearly 7 times longer than that in the dog, and yet the longevity of the horse is not more than 3 times that of the dog. The goat, which takes much longer than the dog to double its weight, has a shorter total life.
I observed myself that new-born mice quadruple their weight in the first 24 hours. The doubling of weight in their case requires a time 36 times less long than that of the cat, and yet the cat lives only 5 times as long as the mouse.
It is fair to say, however, that Bunge himself does not draw a definite conclusion from these figures and has published them only to stimulate interest in the subject. He is against the view of Flourens, and points out that although the multiple 5 is valid for man, it is not so in the case of the horse which finishes its growth in 4 years and yet reaches the age of 40 much less often than human beings attain that of 100 years.
Although it is impossible to admit the existence of exact relations between size and the period of growth on the one side, and longevity on the other, in the mode which Buffon and Flourens have followed, it is none the less true that there is something intrinsic in each kind of animal which sets a definite limit to the length of years it can attain. The purely physiological conditions which determine this limit leave room for a considerable amount of variation in longevity. Duration of life therefore, is a character which can be influenced by the environment. Weismann in his well-known essay on the duration of life, has laid stress on this side of the problem. Longevity, according to him, although in the last resort depending on the physiological properties of the cells of which the organism is composed, can be adapted to the conditions of existence and influenced by natural selection, like other characters useful for the existence of the species.
If a species is to remain in existence, its members must be able to reproduce and the progeny must be able to reach adult life so that they in their turn may reproduce. Now, it happens that there are some animals the fecundity of which is extremely limited. Most birds which are adapted to aerial life, and the weight of which is therefore to be kept down, lay very few eggs. This happens in the case of birds of prey, such as eagles and vultures. These birds nest only once a year, and generally rear two or frequently only a single nestling. In such circumstances the duration of life becomes a factor in the preservation of the species, more important since eggs and chicks are subject to many dangers. Eggs are devoured by many kinds of animals, whilst unseasonable cold may kill the chicks. If the members of such a species were incapable of living long, the unfavourable conditions of life would soon lead to extinction. Those animals which reproduce rapidly generally have a relatively brief duration of life. Mice, rats, rabbits, and many other rodents seldom live more than 5 or 10 years, but reproduce with enormous rapidity. It is almost possible to imagine that there is some sort of intimate link, possibly physiological, between longevity and low fertility. It is a current opinion that reproduction wastes the maternal organism and that mothers of many children grow old prematurely and seldom reach an advanced age. This would seem to mean that fecundity was the cause of the short duration of life. However, we must guard ourselves against such a theory. Longevity, at least in the case of vertebrate animals, differs extremely little in the two sexes, although the cost of the new generation to the adult organism is very much greater in the case of the female than of the male parent. None the less, females frequently reach a great age, especially in the human race where women reach 100 years, or live beyond that time, much more often than men.
Low fertility, however, cannot itself be regarded as a cause of longevity, as there are some very fertile animals which none the less attain great ages. There are parrots which lay two or three times a year, producing six to nine eggs in each clutch. The ducks (Anatidæ) are distinguished for considerable longevity and very high fertility, each nest containing rarely less than six and sometimes as many as sixteen eggs. The common Sheldrake lays from twenty to thirty eggs. Tame ducks, in some parts of the tropics, lay an egg daily throughout the season. Wild ducks lay from seven to fourteen eggs in one nest. Ducks and geese, none the less, frequently attain considerable ages, ducks having been known to live for 29 years. Even the common fowl, which is a notoriously prolific bird, may reach an age of twenty to thirty years.
It will be said, however, that these birds are exposed to many enemies during youth. Chickens, ducklings, and goslings are ready prey for hawks, foxes and small carnivora. The longevity is possibly to be explained as an adaptation for the preservation of the species by compensating for the great destruction of the young. Weismann explains in this way the longevity of many aquatic birds and other creatures that are much preyed on. It must be noted, however, that the longevity cannot depend on the risks run by the young birds, but must have arisen independently. If this had not occurred, creatures, the young of which are destroyed in great numbers, would have ceased to exist, as many species have disappeared in geological time. The longevity of prolific animals, the young of which are destroyed in numbers, must be due to some cause which is neither fertility nor the destruction of their offspring. This cause must be sought in the physiological processes of the organism and can be attributed neither to the length of the period of growth nor to the size attained by the adults.
After having discussed various theories of the cause of the duration of life, M. Oustalet,34 in a most interesting essay on the longevity of vertebrates, came to the conclusion that diet was the chief factor. He thinks that there is a “definite relation between diet and longevity. For the most part herbivorous animals live longer than carnivorous forms, probably because the former find their food with ease and regularity, whilst the latter alternate between semi-starvation and repletion.” There are certainly many instances which give support to the view. Elephants and parrots, for instance, are vegetarian and reach very great ages. On the other hand, there exist long-living carnivorous animals. Many observations have made it certain that owls and eagles reach great ages, and these birds live on animal food. Ravens, which live on carrion, are also notorious for the duration of their lives. There is no exact knowledge as to the ages reached by crocodiles, but although these live on flesh, it is certain that their longevity is great.
We must seek elsewhere for the real factors that control duration of life. Before stating my conclusion, I will review what is known as to the duration of life of different animals.