Essays Upon Heredity and Kindred Biological Problems. Weismann August

further expansions, partly proofs of the views brought forward in the text, and above all a compilation of all the observations which are known to me upon the duration of life in several groups of animals. I am indebted to several eminent specialists for the communication of many data, which are among the most exact that I have been able to obtain. Thus Dr. Hagen of Cambridge (U.S.A.) was kind enough to send me an account of his observations upon insects of different orders: Mr. W. H. Edwards of West Virginia, and Dr. Speyer of Rhoden—their experience with butterflies. Dr. Adler of Schleswig sent me data upon the duration of life in Cynipidae, which have a special value, as they are accompanied by very exact observations upon the conditions of life in these animals; hence in this case we can directly examine the factors upon which, as I believe, the duration of life is chiefly based. Sir John Lubbock in England, and Dr. August Forel of Zürich, have had the kindness to send me an account of their observations upon ants, and S. Clessin of Ochsenfurth his researches upon our native land and fresh-water Mollusca.

      In publishing these valuable communications, together with all facts which I have been able to collect from literature upon the subject of the duration of life, and the little which I have myself observed upon this subject, I hope to provide a stimulus for further observation in this field, which has been hitherto much neglected. The views which I have brought forward in this paper are based on a comparatively small number of facts, at least as far as the duration of life in various species is concerned. The larger the number of accurate data which are supplied, and the more exactly the duration of life and its conditions are ascertained, the more securely will it be possible to establish our views upon the causes which determine the duration of life.

A. W.

      Naples, Dec. 6, 1881.

      I.

      THE DURATION OF LIFE

      With your permission, I will bring before you to-day some thoughts upon the subject of the duration of life. I can scarcely do better than begin with the simple but significant words of Johannes Müller: ‘Organic bodies are perishable; while life maintains the appearance of immortality in the constant succession of similar individuals, the individuals themselves pass away.’

      Omitting, for the time being, any discussion as to the precise accuracy of this statement, it is at any rate obvious that the life of an individual has its natural limit, at least among those animals and plants which are met with in every-day life. But it is equally obvious that the limits are very differently placed in the various species of animals and plants. These differences are so manifest that they have given rise to popular sayings. Thus Jacob Grimm mentions an old German saying, ‘A wren lives three years, a dog three times as long as a wren, a horse three times as long as a dog, and a man three times as long as a horse, that is eighty-one years. A donkey attains three times the age of a man, a wild goose three times that of a donkey, a crow three times that of a wild goose, a deer three times that of a crow, and an oak three times the age of a deer.’

      If this be true a deer would live 6000 years, and an oak nearly 20,000 years. The saying is certainly not founded upon exact observation, but it becomes true if looked upon as a general statement that the duration of life is very different in different organisms.

      The question now arises as to the causes of these great differences. How is it that individuals are endowed with the power of living long in such very various degrees?

      One is at first tempted to seek the answer by an appeal to the differences in morphological and chemical structure which separate species from one another. In fact all attempts to throw light upon the subject which have been made up to the present time lie in this direction.

      All these explanations are nevertheless insufficient. In a certain sense it is true that the causes of the duration of life must be contained in the organism itself, and cannot be found in any of its external conditions or circumstances. But structure and chemical composition—in short the physiological constitution of the body in the ordinary sense of the words—are not the only factors which determine duration of life. This conclusion forces itself upon our attention as soon as the attempt is made to explain existing facts by these factors alone: there must be some other additional cause contained in the organism as an unknown and invisible part of its constitution, a cause which determines the duration of life.

      The size of the organism must in the first place be taken into consideration. Of all organisms in the world, large trees have the longest lives. The Adansonias of the Cape Verd Islands are said to live for 6000 years. The largest animals also attain the greatest age. Thus there is no doubt that whales live for some hundreds of years. Elephants live 200 years, and it would not be difficult to construct a descending series of animals in which the duration of life diminishes in almost exact proportion to the decrease in the size of the body. Thus a horse lives forty years, a blackbird eighteen, a mouse six, and many insects only a few days or weeks.

      If however the facts are examined a little more closely it will be observed that the great age (200 years) reached by an elephant is also attained by many smaller animals, such as the pike and carp. The horse lives forty years, but so does a cat or a toad; and a sea anemone has been known to live for over fifty years. The duration of life in a pig (about twenty years) is the same as that in a crayfish, although the latter does not nearly attain the hundredth part of the weight of a pig.

      It is therefore evident that length of life cannot be determined by the size of the body alone. There is, however, some relation between these two attributes. A large animal lives longer than a small one because it is larger; it would not be able to become even comparatively large unless endowed with a comparatively long duration of life.

      Apart from all other reasons, no one could imagine that the gigantic body of an elephant could be built up like that of a mouse in three weeks, or in a single day like that of the larva of certain flies. The gestation of an elephant lasts for nearly two years, and maturity is only reached after a lapse of about twenty-four years.

      Furthermore, to ensure the preservation of the species, a longer time is required by a large animal than by a small one, when both have reached maturity. Thus Leuckart and later Herbert Spencer have pointed out that the absorbing surface of an animal only increases as the square of its length, while its size increases as the cube; and it therefore follows that the larger an animal becomes, the greater will be the difficulty experienced in assimilating any nourishment over and above that which it requires for its own needs, and therefore the more slowly will it reproduce itself.

      But although it may be stated generally that the duration of the period of growth and length of life are longest in the largest animals, it is nevertheless impossible to maintain that there is any fixed relation between the two; and Flourens was mistaken when he considered that the length of life was always equivalent to five times the duration of the period of growth. Such a conclusion might be accepted in the case of man if we set his period of growth at twenty years and his length of life at a hundred; but it cannot be accepted for the majority of other Mammalia. Thus the horse lives from forty to fifty years, and the latter age is at least as frequently reached among horses as a hundred years among men; but the horse becomes mature in four years, and the length of its life is thus ten or twelve times as long as its period of growth.

      The second factor which influences the duration of life is purely physiological: it is the rate at which the animal lives, the rapidity with which assimilation and the other vital processes take place. Upon this point Lotze remarks in his Microcosmus—‘Active and restless mobility destroys the organized body: the swift-footed animals hunted by man, as also dogs, and even apes, are inferior in length of life to man and the larger beasts of prey, which satisfy their needs by a few vigorous efforts.’ ‘The inertness of the Amphibia is, on the other hand, accompanied by relatively great length of life.’

      There is certainly some truth in these observations, and yet it would be a great mistake to assume that activity necessarily implies a short life. The most active birds have very long lives, as will be shown later on: they live as long as and sometimes longer than the majority of Amphibia which reach the same size. The organism must not be looked upon as a heap of combustible material, which is completely reduced to ashes in a certain time the length of which is determined by size, and by the rate at which it burns; but it should be rather compared to a fire, to which fresh fuel can be continually