LONGEVITY, a term applied to express either the length or the duration of life in any organism, but, as cases of long duration excite most interest, frequently used to denote a relatively unusual prolongation of life. There is no reason to suppose that protoplasm, the living material of organisms, has a necessarily limited duration of life, provided that the conditions proper to it are maintained, and it has been argued that since every living organism comes into existence as a piece of the protoplasm of a pre-existing living organism, protoplasm is potentially immortal. Living organisms exist, however, as particles or communities of particles of protoplasm (see Life), and as such have a limited duration of life. Longevity, as E. Ray Lankester pointed out in 1869, for practical purposes must be understood to mean the “length of time during which life is exhibited in an individual.” The word “individual” must be taken in its ordinary sense as a wholly or partially independent, organized mass produced from a pre-existing organized mass, as otherwise the problem will be confused by arguments as to the meaning of biological individuality.
Empirical Data.—A multitude of observations show that only a very brief life, ranging from a few hours to a few days, is the normal fate of the vast majority of single-celled organisms, whether these be animal or vegetable or on the border-line between the two kingdoms. Death comes to them rapidly from internal or external causes, or the individual life ends in conjugation or division or spore-formation. Under special conditions, natural or artificial, the individual life may be prolonged by desiccation, or freezing, or by some similar arrest of functional activity.
The duration of life among plants is varied. The popular division into annuals, biennials and perennials is not absolute, for natural and artificial conditions readily prolong the lives of annuals and biennials for several seasons, whereas the case of perennials is much complicated by the mode of growth, and the problem of individuality, however we desire to exclude it, obtrudes itself. In the vast majority of cases where a plant is obviously a simple individual, its life is short, ranging from a few days in the case of fungi, to two seasons in the case of biennial herbs. Most of the simple algae are annual, their life enduring only for part of the year; the branching algae are more often perennial, but in their cases not only are observations as to duration lacking, but however simply we may use the term individual, its application is difficult. The larger terrestrial plants with woody tissues which we denote roughly as shrubs and trees have an individuality which, although different from that of a hyacinth or carrot, is usually obvious. Shrubs live from four to ten or more years, and it apparently is the case that odoriferous shrubs such as sage and lavender display the longer duration. Trees with soft wood, such as poplars and willows, last for about fifty years, fruit-trees rather longer. Estimates of the age which large trees can attain, based partly on attempts to count the annual rings, have been given by many writers, and range from about three hundred years in the case of the elm to three to five thousand years in the case of Sequoia gigantea of California, and over five thousand years in that of the baobab (Adansonia digitata) of Cape Verde. It is impossible to place exact reliance on these estimates, but it is at least certain that very many trees have a duration of life exceedingly great in comparison with the longest-lived animals.
The duration of life amongst multicellular invertebrate animals is little known, except in the frequent instances where it is normally brief. Many sponges and polyps die at the end of the season, leaving winter eggs or buds. The much-branched masses of the larger sponges and compound hydrozoa certainly may be perennial. A sea-anemone (Actinia mesembryanthemum), captured in 1828 by Sir John Dalyell, a Scottish naturalist, and then guessed to be about seven years old, lived in captivity in Edinburgh until 1887, the cause of death being unknown. As other instances of great ages attained by sea-anemones are on record, it is plain that these animals, although simple polyps, are long-lived. Echinoderms are inferred to live to considerable ages, as they grow slowly and as there is great difference in size amongst fully adult specimens. On similar reasoning, considerable age is attributed to the larger annulates and crustacea, but the smaller forms in many cases are known to have very short lives. The variation in the length of life of molluscs appears to be great. Many species of gastropods live only a few years; others, such as Natica heros, have reached thirty years, whilst the large Tridacna gigas is stated to live from sixty to a hundred years. Among insects, the adult stage has usually only a very short duration of life, extending from a few hours to a few months, but the larval stages may last much longer. Including these latter, the range of duration among insects, taking the whole life from hatching to death, appears to lie between the limits of a few weeks in the case of plant-lice to seventeen years in the case of the American Cicada septemdecim, the larva of which lives seventeen years, the adult only a month. Most butterflies are annuals, but those which fail to copulate may hibernate and live through a second season, whilst the lives of some have been preserved artificially for seven years. Worker bees and drones do not survive the season, but queens may live from two to five years. In the case of vertebrates, the duration of life appears to be greater among fish and reptiles than among birds and mammals. The ancient Romans have noted that eels, kept in aquaria, could reach the age of sixty years. Estimates based on size and rate of growth have led to the inference that salmon may live to the age of a hundred years, whilst G. L. L. Buffon set down the period of life of carp in ponds as one hundred and fifty years, and there is evidence for a pike having reached the age of over two centuries. More recently it has been claimed that the age of fish can be ascertained exactly by counting the annual rings of the otoliths. No great ages have as yet been recorded by this method, whilst, on the other hand, by revealing great variations of weight and size in fishes with the same number of annual rings, it has thrown doubt on the validity of estimates of age based on size and rate of growth. The evidence as a whole is unsatisfactory, but it is highly probable that in the absence of accidents most fish can attain very great ages. The duration of life among batrachia is little known, but small frogs have been recorded as living over twelve years, and toads up to thirty-six years.
Almost nothing is known as to the longevity of snakes and lizards, but it is probable that no great ages are reached. Crocodiles, alligators and caymans grow slowly and are believed to live very long. There is exact evidence as to alligators in captivity in Europe reaching forty years without signs of senescence, and some of the sacred crocodiles of India are believed to be more than a hundred years old. Chelonians live still longer. A tortoise has lived for eighty years in the garden of the governor of Cape Town, and is believed to be at least two hundred years old. There are records of small land-tortoises that have been kept in captivity for over a century, whilst the very large tortoises of the Galapagos Islands certainly attain ages of at least two centuries and possibly much more. A considerable body of information exists regarding the longevity of birds, and much of this has been brought together by J. H. Gurney. From his lists, which include more than fifty species, it appears that the duration is least in the case of small passerine and picarian birds, where it ranges from eight or nine years (goat-suckers and swifts) to a maximum of twenty-five years, the latter age having been approached by larks, canaries and goldfinch. Gulls have been recorded as living over forty years, ducks and geese over fifty years (the duchess of Bedford has recorded the case of a Chinese goose having been in possession of the same family for fifty-seven years). Parrots frequently live over eighty years, swans nearly as long, ravens and owls rather less, whilst there is excellent evidence of eagles and falcons considerably exceeding a hundred years. Notwithstanding their relatively large size, struthious birds do not reach great ages. The records for cassowaries and rheas do not exceed thirty years, and the maximum for ostriches is fifty years, and that on doubtful evidence.
Exact records regarding the longevity of mammals are surprisingly few. There is no evidence as to Monotremes. The life of Marsupials in captivity is seldom long; a phalanger has lived in the London Zoological Gardens and showed no signs of age at more than ten years old; it may be inferred that the larger forms are capable of living longer. Reliable records as to Edentates do not exist; those in captivity have short lives, but the size and structure of some of the extinct forms suggests that they may have reached a great age. Nothing is known regarding the longevity of Sirenians, except that they do not live long in captivity. In the case of Cetaceans, estimates based on the growth of whale-bone assign an age of several centuries to whale-bone whales; exact records do not exist. More is known regarding Ungulates, as many of these are domesticated, semi-domesticated or are frequently kept in captivity. Great length of life has been assigned to the rhinoceros, but the longest actual record is that of an Indian rhinoceros which lived for thirty-seven years in the London Zoological Gardens. The usual duration of life in the case of horses, asses and zebras is from fifteen to thirty years, but instances of individuals reaching fifty years are fairly well authenticated. Domestic cattle may live from twenty-five to thirty years, sheep and goats from twelve to fourteen years, antelopes rather longer, especially in the case of the larger forms. A giraffe has lived for nineteen years in the London Zoological Gardens. Deer are reputed to live longer than sheep, and records of individuals at the London Gardens confirm this, but it is doubtful if they live as long as cattle. Camels are long-lived, according to repute, but actual records show no great age; a llama which died in the London Gardens at the age of seventeen years showed unmistakable signs of senility. The hippopotamus is another large ungulate to which great longevity has been assigned, but the longest actual record is the case of a female born in the London Gardens which died in its thirty-fifth year. The duration of life assigned to domestic swine is about twenty years; an Indian wild boar, alive in the London Zoological Gardens in 1910, and apparently in full vigour, was fifteen years old. Elephants are usually supposed capable of reaching great ages, but the actual records of menagerie and military animals show that thirty to forty years is a normal limit. Facts as to rodents are not numerous; the larger forms such as hares and rabbits may live for ten years, smaller forms such as rats and mice, for five or six years. Bats have a reputation for long duration of life, and tropical fruit-bats are known to have lived for seventeen years. No great ages have been recorded for Carnivora, but the average is fairly high. Twenty-five years appears to be a limit very rarely exceeded by lions, tigers or bears; domestic cats may live for from twelve to twenty-three years, and dogs from sixteen to eighteen years, though cases of as many as thirty-four years have been noted. Less is known of the smaller forms, but menagerie records show that ages between twelve and twenty are frequently reached. There were in 1910 in the London Zoological Gardens, apparently in good health, a meerkat at least twelve years old, a sand-badger fourteen years and a ratel nineteen years of age. Records regarding monkeys are unsatisfactory, for these creatures are notoriously delicate in captivity, and it is practically certain that under such circumstances they rarely die of old age. A grey lemur eleven years old and a chimpanzee eleven and a half, both in good health in the London Zoological Gardens, appear to be the oldest primates definitely recorded. Estimates based on size, condition of the skull and so forth obtained by examination of wild specimens that have been killed would seem to establish a rough correspondence between the size of monkeys and their duration of life, and to set the limits as between seven or eight and thirty years.
With regard to the human race, there seems to be almost no doubt but that the average duration of life has increased with civilization; the generally improved conditions of life, the greater care of the young and of the aged and the advance in medical and surgical science far more than outweigh any depressing effect caused by the more strenuous and nervous activity required by modern social organization. The expectation of life of those who attain the age of sixty varies with race, sex and occupation, but is certainly increasing, and an increasing number of persons have a chance of reaching and do reach ages between ninety and one hundred. Careful investigation has thrown doubt almost amounting to disproof on the much-quoted cases of great longevity, such as that of Thomas Parr, the Shropshire peasant, who was supposed to have reached his hundred and fifty-third year, and, although the existence of centenarians is thoroughly established, any ages exceeding a hundred by more than two or three years are, at the most, dubious.
A survey of the facts of longevity, so far as these are established on reasonable evidence, discloses that the recorded ages both of men and animals are much shorter than those assigned in popular belief. The duration of life is usually brief in the animal kingdom, and except for some fish and reptiles, and possibly whales, it is certain that a man enjoys the longest average duration of life and that centenarians occur more frequently amongst men than amongst most of the lower animals.
Theories of Longevity.—Ray Lankester has pointed out that several meanings are attached to the word longevity. It may be used of an individual, and in this sense has little importance, partly because of the inevitable variability of the individual, and partly because there may be individuals that are abnormal in duration of life, just as there are abnormalities in weight or height. It may be used for the average duration of life of all the individuals of a species and so be another way of expressing the average mortality that affects the species, and that varies not only with structure and constitution but with the kind of enemies, accidents and conditions to which the members of the species are subject. If we reflect on the large incidence of mortality from external causes affecting a species and particularly the young of a species, we shall see that we must conclude that intrinsic, physiological causes can have relatively little weight in determining the average mortality rate. Finally, longevity may be used, and is most conveniently used, to denote the specific potential longevity, that is to say the duration of life that would be attained by normal individuals of a species if the conditions were most favourable. It is necessary to keep in mind these various applications of the term when considering the theoretical explanations that have been associated with the empirical facts.
There is a certain relation between size and longevity. As a general rule small animals do not live so long as larger creatures. Whales survive elephants, elephants live longer than camels, horses and deer, and these again than rabbits and mice. But the relation is not absolute; parrots, ravens and geese live longer than most mammals and than many larger birds. G. L. L. Buffon tried to find a more definite measure of longevity, and believed that it was given by the ratio between the whole period of life and the period of growth. He believed that the possible duration of life was six or seven times that of the period of growth. Man, he said, takes fourteen years to grow, and his duration of life is ninety to one hundred years; the horse has reached its full size at four years of age and may live for a total period of twenty-five to thirty years. M. J. P. Flourens attempted to make Buffon’s suggestion more exact; he took the end of the period of growth as the time at which the epiphyses of the long bones united with the bones themselves, and on this basis held that the duration of life was five times the length of the period of growth. The theories of Buffon and Flourens, however, do not apply to all vertebrates and have no meaning in the case of invertebrates. Y. Bunge has suggested that in the case of mammals the period taken by the new-born young to double in weight is an index of the rapidity of growth and is in a definite relation to the possible duration of life. M. Oustalet has discussed the existence of definite relations between duration of life and size, rate of growth, period of gestation and so forth, and found so many exceptions that no general conclusion could be drawn. He finally suggested that diet was the chief factor in determining the span of life. E. Metchnikoff has provided the most recent and fullest criticism and theory of the physiological causes of longevity. He admits that many factors must be involved, as the results vary so much in different kinds of animals. He thinks that too little is known of the physiological processes of invertebrates to draw any valid conclusions in their case. With regard to vertebrates, he calls attention to the gradual reduction of longevity as the scale of life is ascended. On the whole, reptiles live much longer than birds, and birds than mammals, the contrast being specially notable when birds and mammals are compared. He dismisses the effect of the reproductive tax from possible causes of short duration of life, for the obvious reason that longevity is nearly equal in the two sexes, although females have a much greater reproductive drain. He points out that the hind-gut or large intestine is least developed in fishes, relatively small in reptiles, still small but relatively larger in birds and largest in mammals, relatively and absolutely, the caecum or caeca being reckoned as part of the hind-gut. The area of the intestinal tract in question is of relatively little importance in digestion, although a considerable amount of absorption may take place from it. It serves chiefly as a reservoir of waste matter and is usually the seat of extensive putrefactive change. The products of putrefaction are absorbed by the blood and there results a constant auto-intoxication of the body which Metchnikoff believes to be the principal agent in senile degeneration. Mammals, if they escape from enemies, diseases and accidents, fall victims to premature senility as the result of the putrefactive changes in their intestines, and the average mortality of the species is much too high, the normal specific longevity being rarely if ever attained. Metchnikoff urges, and so far probably is followed by all competent authorities, that improvements in the conditions of life, greater knowledge of disease and of hygiene and simplification of habits are tending to reduce the average mortality of man and the domestic animals, and to bring the average longevity nearer the specific longevity. He adds to this, however, a more special theory, which, although it appears rapidly to be gaining ground, is yet far from being accepted. The theory is that duration of life may be prolonged by measures directed against intestinal putrefaction.
The process of putrefaction takes place in masses of badly-digested food, and may be combated by careful dieting, avoidance of rich foods of all kinds and particularly of flesh and alcohol. Putrefaction, however, cannot take place except in the presence of a particular group of bacteria, the entrance of which to the body can be prevented to a certain extent. But it would be impossible or impracticable to secure a sterilized diet, and Metchnikoff urges that the bacteria of putrefaction can be replaced or suppressed by another set of microbes. He found that there was a widely spread popular belief in the advantage of diet consisting largely of products of soured milk and that there was a fair parallel between unusual longevity and such a diet. Experimentally he showed that the presence of the bacilli which produce lactic acid inhibited the process of putrefaction. Accordingly he recommends that the diet of human beings should include preparations of milk soured by cultures of selected lactic acid bacilli, or that the spores of such bacilli should be taken along with food favourable to their development. In a short time the bacilli establish themselves in the large intestine and rapidly stop putrefactive change. The treatment has not yet been persisted in sufficiently long by a sufficient number of different persons to be accepted as universally satisfactory, and there is even more difference of opinion as to Metchnikoff’s theory that the chief agent in senile degeneration is the stimulation of phagocytes by the products of putrefaction with the resulting destruction of the specific cells of the tissues. Metchnikoff, however, gave it to the world, not as a proved and completed doctrine, but as the line of inquiry that he himself had found most promising. He has suggested further that if the normal specific longevity were attained by human beings, old and not degenerate individuals would lose the instinct for life and acquire an instinct for death, and that as they had fulfilled the normal cycle of life, they would accept death with the same relieved acquiescence that they now accept sleep.
The various writers whose opinions have been briefly discussed agree in supposing that there is a normal specific longevity, although Metchnikoff alone has urged that this differs markedly from the average longevity, and has propounded a theory of the causes of the divergence. It is common ground that they believe the organism to be wound up, so to say, for a definite period, but have no very definite theory as to how this period is determined. A. Weismann, on the other hand, in a well-known essay on the duration of life, has developed a theory to explain the various fashions in which the gift of life is measured out to different kinds of creatures. He accepts the position that purely physiological conditions set a limit to the number of years that can be attained by each kind of multi-cellular organism, but holds that these conditions leave room for a considerable amount of variation. Duration of life, in fact, according to Weismann, is a character that can be influenced by the environment and that by a process of natural selection can be adapted to the conditions of existence of different species.
If a species is to maintain its existence or to increase, it is obvious that its members must be able to replace the losses caused by death. It is necessary, moreover, for the success of the species, that an average population of full vigour should be maintained. Weismann argues that death itself is an adaptation to secure the removal of useless and worn-out individuals and that it comes as soon as may be after the period of reproductive activity. It is understood that the term reproductive activity covers not merely the production of new individuals but the care of these by the parents until they are self-sufficient. The average longevity, according to Weismann, is adapted to the needs of the species; it is sufficiently long to secure that the requisite number of new individuals is produced and protected. He has brought together a large number of instances which show that there is a relation between duration of life and fertility. Birds of prey, which breed slowly, usually producing an annual brood of no more than one or two, live to great ages, whilst rabbits which produce large litters at frequent intervals have relatively short lives. Allowance has to be made in cases where the young are largely preyed upon by enemies, for this counteracts the effect of high fecundity. In short, the duration of life is so adapted that a pair of individuals on the average succeed in rearing a pair of offspring. Metchnikoff, however, has pointed out that the longevity of such fecund creatures must have arisen independently, as otherwise species subject to high risks of this nature would have ceased to exist and would have disappeared, as many species have vanished in the past of the world’s history.
The normal specific longevity, the age to which all normal individuals of a species would survive under the most favourable conditions, must depend on constitution and structure. No doubt selection is involved, as it is obvious that creatures would perish if their constitution and structure were not such that they could live long enough to reproduce their kind. The direct explanation, however, must be sought for in size, complexity of structure, length of period of growth, capacity to withstand the wear and tear of life and such other intrinsic qualities. The average specific longevity, on the other hand, depends on a multitude of extrinsic conditions operating on the intrinsic constitution; these extrinsic conditions are given by the environment of the species as it affects the young and the adults, enemies, diseases, abundance of food, climatic conditions and so forth. It would seem most natural to suppose that in all cases, except perhaps those of intelligent man and the domestic animals or plants he harbours, the average longevity must vary enormously with changing conditions, and must be a factor of greater importance in the survival of the species than the ideal normal specific longevity. It also seems more probable that the reproductive capacity, which is extremely variable, has been adapted to the average longevity of the species, than that, as Weismann supposed, it should itself be the determining cause of the duration of life.
References.—G. L. L. Buffon, Histoire naturelle générale et particulière, vol. ii. (Paris, 1749); Y. Bunge, Archiv. f. die gesammte Physiologie, vol. xcv. (Bonn, 1903); M. J. P. Flourens, De la longévité humaine et de la quantité de vie sur le globe (Paris, 1855); J. H. Gurney, On the Comparative Ages to which Birds live, Ibis, p. 19 (1899); Sir E. Ray Lankester, Comparative Longevity in Man and the Lower Animals (London, 1870); E. Metchnikoff, The Prolongation of Life (London, 1908); M. Oustalet, La Nature, p. 378 (1900); A. Weismann, Essays upon Heredity (Oxford, 1889). (P. C. M.)