Essays Upon Heredity and Kindred Biological Problems

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 added, and which, whether it burns quickly or slowly, can be kept burning as long as necessity demands.

The connection between activity and shortness of life cannot be explained by supposing that a more rapid consumption of the body occurs, but it is explicable because the increased rate at which the vital processes take place permit the more rapid achievement of the aim and purpose of life, viz. the attainment of maturity and the reproduction of the species.

When I speak of the aim and purpose of life, I am only using figures of speech, and I do not mean to imply that nature is in any way working consciously.

When I was speaking of the relation between duration of life and the size of the body, I might have added another factor which also exerts some influence, viz. the complexity of the structure. Two organisms of the same size, but belonging to different grades of organization, will require different periods of time for their development. Certain animals of a very lowly organization, such as the Rhizopoda, may attain a diameter of ·5 mm. and may thus become larger than many insects’ eggs. Yet under favourable circumstances an Amoeba can divide into two animals in ten minutes, while no insect’s egg can develope into the young animal in a less period than twenty-four hours. Time is required for the development of the immense number of cells which must in the latter case arise from the single egg-cell.

Hence we may say that the peculiar constitution of an animal does in part determine the length of time which must elapse before reproduction begins. The period before reproduction is however only part of the whole life of an animal, which of course extends over the total period during which the animal exists.

Hitherto it has always been assumed that the duration of this total period is solely determined by the constitution of the animal’s body. But the assumption is erroneous. The strength of the spring which drives the wheel of life does not solely depend upon the size of the wheel itself or upon the material of which it is made; and, leaving the metaphor, duration of life is not exclusively determined by the size of the animal, the complexity of its structure, and the rate of its metabolism. The facts are plainly and clearly opposed to such a supposition.

How, for instance, can we explain from this point of view the fact that the queen-ant and the workers live for many years, while the males live for a few weeks at most? The sexes are not distinguished by any great difference in size or complexity of body, or in the rate of metabolism. In all these three particulars they must be looked upon as precisely the same, and yet there is this immense difference between the lengths of their lives.

I shall return later on to this and other similar cases, and for the present I assume it to be proved that physiological considerations alone cannot determine the duration of life. It is not these which alone determine the strength of the spring which moves the machinery of life; we know that springs of different strengths may be fixed in machines of the same kind and quality. This metaphor is however imperfect, because we cannot imagine the existence of any special force in an organism which determines the duration of its life; but it is nevertheless useful because it emphasises the fact that the duration of life is forced upon the organism by causes outside itself, just as the spring is fixed in its place by forces outside the machine, and not only fixed in its place, but chosen of a certain strength so that it will run down after a certain time.

To put it briefly, I consider that duration of life is really dependent upon adaptation to external conditions, that its length, whether longer or shorter, is governed by the needs of the species, and that it is determined by precisely the same mechanical process of regulation as that by which the structure and functions of an organism are adapted to its environment.

Assuming for the moment that these conclusions are valid, let us ask how the duration of life of any given species can have been determined by their means. In the first place, in regulating duration of life, the advantage to the species, and not to the individual, is alone of any importance. This must be obvious to any one who has once thoroughly thought out the process of natural selection. It is of no importance to the species whether the individual lives longer or shorter, but it is of importance that the individual should be enabled to do its work towards the maintenance of the species. This work is reproduction, or the formation of a sufficient number of new individuals to compensate the species for those which die. As soon as the individual has performed its share in this work of compensation, it ceases to be of any value to the species, it has fulfilled its duty and may die. But the individual may be of advantage to the species for a longer period if it not only produces offspring, but tends them for a longer or shorter time, either by protecting, feeding, or instructing them. This last duty is not only undertaken by man, but also by animals, although to a smaller extent; for instance, birds teach their young to fly, and so on.

We should therefore expect to find that, as a rule, life does not greatly outlast the period of reproduction except in those species which tend their young; and as a matter of fact we find that this is the case.

All mammals and birds outlive the period of reproduction, but this never occurs among insects except in those species which tend their young. Furthermore, the life of all the lower animals ceases also with the end of the reproductive period, as far as we can judge.

Duration of life is not however determined in this way, but only the point at which its termination occurs relatively to the cessation of reproduction. The duration itself depends first upon the length of time which is required for the animal to reach maturity—that is, the duration of its youth, and, secondly, upon the length of the period of fertility—that is the time which is necessary for the individual to produce a sufficient number of descendants to ensure the perpetuation of the species. It is precisely this latter point which is determined by external conditions.

There is no species of animal which is not exposed to destruction through various accidental agencies—by hunger or cold, by drought or flood, by epidemics, or by enemies, whether beasts of prey or parasites. We also know that these causes of death are only apparently accidental, or at least that they can only be called accidental as far as a single individual is concerned. As a matter of fact a far greater number of individuals perish through the operation of these agencies than by natural death. There are thousands of species of which the existence depends upon the destruction of other species; as, for example, the various kinds of fish which feed on the countless minute Crustacea inhabiting our lakes.

It is easy to see that an individual is, ceteris paribus, more exposed to accidental death when the natural term of its life becomes longer; and therefore the longer the time required by an individual for the production of a sufficient number of descendants to ensure the existence of the species, the greater will be the number of individuals which perish accidentally before they have fulfilled this important duty. Hence it follows, first, that the number of descendants produced by any individual must be greater as the duration of its reproductive period becomes longer; and, secondly, the surprising result that nature does not tend to secure the longest possible life to the adult individual, but, on the contrary, tends to shorten the period of reproductive activity as far as possible, and with this the duration of life; but these conclusions only refer to the animal and not to the vegetable world.

All this sounds very paradoxical, but the facts show that it is true. At first sight numerous instances of remarkably long life seem to refute the argument, but the contradictions are only apparent and disappear on closer investigation.

Birds as a rule live to a surprisingly great age. Even the smallest of our native singing birds lives for ten years, while the nightingale and blackbird live from twelve to eighteen years. A pair of eider ducks were observed to make their nest in the same place for twenty years, and it is believed that these birds sometimes reach the age of nearly one hundred years. A cuckoo, which was recognised by a peculiar note in its call, was heard in the same forest for thirty-two consecutive years. Birds of prey, and birds which live in marshy districts, become much older, for they outlive more than one generation of men.

Schinz mentions a bearded vulture which was seen sitting on a rock upon a glacier near Grindelwald, and the oldest men in Grindelwald had, when boys, seen the same bird sitting on the same rock. A white-headed vulture in the Schönbrunn Zoological Gardens had been in captivity for 118 years, and many examples are known of eagles and falcons reaching an age of over 100 years. Finally, we must not forget Humboldt’s[1 - Humboldt’s ‘Ausichten der Natur.’] Atur parrot from the Orinoco, concerning which the Indians said that it could not be understood because it spoke the language of an extinct tribe.

It is therefore necessary to ask how far we can show that such long lives are really the shortest which are possible under the circumstances.

Two factors must here be taken into consideration; first, that the young of birds are greatly exposed to destructive agencies; and, secondly, that the structure of a bird is adapted for flight and therefore excludes the possibility of any great degree of fertility.

Many birds, like the stormy petrel, the diver, guillemot, and other sea-birds, lay only a single egg, and breed (as is usually the case with birds) only once a year. Others, such as birds of prey, pigeons, and humming-birds, lay two eggs, and it is only those which fly badly, such as jungle fowls and pheasants, which produce a number of eggs (about twenty), and the young of these very species are especially exposed to those dangers which more or less affect the offspring of all birds. Even the eggs of our most powerful native bird of prey, the golden eagle, which all animals fear, and of which the eyrie, perched on a rocky height, is beyond the reach of any enemies, are very frequently destroyed by late frosts or snow in spring, and, at the end of the year in winter, the young birds encounter the fiercest of foes, viz. hunger. In the majority of birds, the egg, as soon as it is laid, becomes exposed to the attacks of enemies; martens and weasels, cats and owls, buzzards and crows are all on the look out for it. At a later period the same enemies destroy numbers of the helpless young, and in winter many succumb in the struggle against cold and hunger, or to the numerous dangers which attend migration over land and sea, dangers which decimate the young birds.

It is impossible directly to ascertain the exact number which are thus destroyed; but we can arrive at an estimate by an indirect method. If we agree with Darwin and Wallace in believing that in most species a certain degree of constancy is maintained in the number of individuals of successive generations, and that therefore the number of individuals within the same area remains tolerably uniform for a certain period of time; it follows that, if we know the fertility and the average duration of life of a species, we can calculate the number of those which perish before reaching maturity. Unfortunately the average length of life is hardly known with certainty in the case of any species of bird. Let us however assume, for the sake of argument, that the individuals of a certain species live for ten years, and that they lay twenty eggs in each year; then of the 200 eggs which are laid during the ten years, which constitute the lifetime of an individual, 198 must be destroyed, and only two will reach maturity, if the number of individuals in the species is to remain constant. Or to take a concrete example; let us fix the duration of life in the golden eagle at 60 years, and its period of immaturity (of which the length is not exactly known) at ten years, and let us assume that it lays two eggs a year;—then a pair will produce 100 eggs in 50 years, and of these only two will develope into adult birds; and thus on an average a pair of eagles will only succeed in bringing a pair of young to maturity once in fifty years. And so far from being an exaggeration, this calculation rather under-estimates the proportion of mortality among the young; it is sufficient however to enforce the fact that the number of young destroyed must reach in birds a very high figure as compared with the number of those which survive [See Note 1 (#x8_pgepubid00008)].

If this argument holds, and at the same time the fertility from physical and other grounds cannot be increased, it follows that a relatively long life is the only means by which the maintenance of the species of birds can be secured. Hence a great length of life is proved to be an absolute necessity for birds.

I have already mentioned that these animals demonstrate most clearly that physiological considerations do not by any means suffice to explain the duration of life. Although all vital processes take place with greater rapidity and the temperature of the blood is higher in birds than in mammals, yet the former greatly surpass the latter in length of life. Only in the largest Mammalia,—the whales and the elephants—is the duration of life equal to or perhaps greater than that of the longest lived birds. If we compare the relative weights of these animals, the Mammalia are everywhere at a disadvantage. Even such large animals as the horse and bear only attain an age of fifty years at the outside; the lion lives about thirty-five years, the wild boar twenty-five, the sheep fifteen, the fox fourteen, the hare ten, the squirrel and the mouse six years [See Note 2 (#x8_pgepubid00009)]; but the golden eagle, though it does not weigh more than from 9-12 pounds, and is thus intermediate as regards weight between the hare and the fox, attains nevertheless an age which is ten times as long. The explanation of this difference is to be found first in the much greater fertility of the smaller Mammalia, such as the rabbit or mouse, and secondly in the much lower mortality among the young of the larger Mammalia. The minimum duration of life necessary for the maintenance of the species is therefore much lower than it is among birds. Even here, however, we are not yet in possession of exact statistics indicating the number of young destroyed; but it is obvious that Mammalia possess over birds a great advantage in their intra-uterine development. In Mammalia the destruction of young only begins after birth, while in birds it begins during the development of the embryo. This distinction is in fact carried even further, for many mammals protect their young against enemies for a long time after birth.

It is unnecessary to go further into the details of these cases, or to consider whether and to what extent every class of the animal kingdom conforms to these principles. Thus to consider all or even most of the classes of the animal kingdom would be quite impossible at the present time, because our knowledge of the duration of life among animals is very incomplete. Biological problems have for a long time excited less interest than morphological ones. There is nothing or almost nothing to be found in existing zoological text books upon the duration of life in animals; and even monographs upon single classes, such as the Amphibia, reptiles, or even birds, contain very little on this subject. When we come to the lower animals, knowledge on this point is almost entirely wanting. I have not been able to find a single reference to the age in Echinodermata, and very little about that of worms, Crustacea, and Coelenterata [See Note 4 (#x8_pgepubid00011)]. The length of life in many molluscan species is very well known, because the age can be determined by markings on the shell [See Note 5 (#x9_pgepubid00012)]. But even in this group, any exact knowledge, such as would be available for our purpose, is still wanting concerning such necessary points as the degree of fertility, the relation to other animals, and many other factors.

Data the most exact in all respects are found among the insects [See Note 3 (#x8_pgepubid00010)], and to this class I will for a short time direct your special attention. We will first consider the duration of larval life. This varies very greatly, and chiefly depends upon the nature of the food, and the ease or difficulty with which it can be procured. The larvae of bees reach the pupal stage in five to six days; but it is well known that they are fed with substances of high nutritive value (honey and pollen), and that they require no great effort to obtain the food, which lies heaped up around them. The larval life in many Ichneumonidae is but little longer, being passed in a parasitic condition within other insects; abundance of accessible food is thus supplied by the tissues and juices of the host. Again, the larvae of the blow-fly become pupae in eight to ten days, although they move actively in boring their way under the skin and into the tissues of the dead animals upon which they live. The life of the leaf-eating caterpillars of butterflies and moths lasts for six weeks or longer, corresponding to the lower nutritive value of their food and the greater expenditure of muscular energy in obtaining it. Those caterpillars which live upon wood, such as Cossus ligniperda, have a larval life of two to three years, and the same is true of hymenopterous insects with similar habits, such as Sirex.

Furthermore, predaceous larvae require a long period for attaining their full size, for they can only obtain their prey at rare intervals and by the expenditure of considerable energy. Thus among the dragon-flies larval life lasts for a year, and among many may-flies even two or three years.

All these results can be easily understood from well-known physiological principles, and they indicate that the length of larval life is very elastic, and can be extended as circumstances demand; for otherwise carnivorous and wood-eating larvae could not have survived in the phyletic development of insects. Now it would be a great mistake to suppose that there is any reciprocal relation between duration of life in the larva and in the mature insect, or imago; or, to put it differently, to suppose that the total duration of life is the same in insects of the same size and activity, so that the time which is spent in the larval state is, as it were, deducted from the life of the imago, and vice versa. That this cannot be the case is shown by the fact already alluded to, that among bees and ants larval life is of the same length in males and females, while there is a difference of some years between the lengths of their lives as imagos.

The life of the imago is generally very short, and not only ends with the close of the period of reproduction, as was mentioned above, but this latter period is also itself extremely short [See Note 3 (#x8_pgepubid00010)].

The larva of the cockchafer devours the roots of plants for a period of four years, but the mature insect with its more complex structure endures for a comparatively short time; for the beetle itself dies in about a month after completing its metamorphosis. And this is by no means an extreme case. Most butterflies have an even shorter life, and among the moths there are many species (as in the Psychidae) which only live for a few days, while others again, which reproduce by the parthenogenetic method, only live for twenty-four hours. The shortest life is found in the imagos of certain may-flies, which only live four to five hours. They emerge from the pupa-case towards the evening, and as soon as their wings have hardened, they begin to fly, and pair with one another. Then they hover over the water; their eggs are extruded all at once, and death follows almost immediately.

The short life of the imago in insects is easily explained by the principles set forth above. Insects belong to the number of those animals which, even in their mature state, are very liable to be destroyed by others which are dependent upon them for food; but they are at the same time among the most fertile of animals, and often produce an astonishing number of eggs in a very short time. And no better arrangement for the maintenance of the species under such circumstances can be imagined than that supplied by diminishing the duration of life, and simultaneously increasing the rapidity of reproduction.

This general tendency is developed to very different degrees according to conditions peculiar to each species. The shortening of the period of reproduction, and the duration of life to the greatest extent which is possible, depends upon a number of co-operating circumstances, which it is impossible to enumerate completely. Even the manner in which the eggs are laid may have an important effect. If the larva of the may-fly lived upon some rare and widely distributed food-plant instead of at the bottom of streams, the imagos would be compelled to live longer, for they would be obliged—like many moths and butterflies—to lay their eggs singly or in small clusters, over a large area. This would require both time and strength, and they could not retain the rudimentary mouth which they now possess, for they would have to feed in order to acquire sufficient strength for long flights; and—whether they were carnivorous like dragon-flies, or honey-eating like butterflies—their feeding would itself cause a further expenditure of both time and strength, which would necessitate a still further increase in the duration of life. And as a matter of fact we find that dragon-flies and swift-flying hawk-moths often live for six or eight weeks and sometimes longer.

We must also remember that in many species the eggs are not mature immediately after the close of the pupal stage, but that they only gradually ripen during the life of the imago, and frequently, as in many beetles and butterflies, do not ripen simultaneously, but only a certain number at a time. This depends, first, upon the amount of reserve nutriment accumulated in the body of the insect during larval life; secondly, upon various but entirely different circumstances, such as the power of flight. Insects which fly swiftly and are continually on the wing, like hawk-moths and dragon-flies, cannot be burdened with a very large number of ripe eggs. In these cases the gradual ripening of the eggs becomes necessary, and involves an increase in the duration of life. In Lepidoptera, we see how the power of flight diminishes step by step as soon as other circumstances permit, and simultaneously how the eggs ripen more and more rapidly, while the length of life becomes shorter, until a minimum is reached. Only two stages in the process of transformation can be mentioned here.

The strongest flyers—the hawk-moths and butterflies—must be looked upon as the most specialised and highest types among the Lepidoptera. Not only do they possess organs for flight in their most perfect form, but also organs for feeding—the characteristic spiral proboscis or ‘tongue.’

There are certain moths (among the Bombyces) of which the males fly as well as the hawk-moths, while the females are unable to use their large wings for flight, because the body is too heavily weighted by a mass of eggs, all of which reach maturity at the same time. Such species, as for instance Aglia tau, are unable to distribute their eggs over a wide area, but are obliged to lay them all in a single spot. They can however do this without harm to the species, because their caterpillars live upon forest trees, which provide abundant food for a larger number of larvae than can be produced by the eggs of a single female. The eggs of Aglia tau are deposited directly after pairing, and shortly afterwards the insect dies at the foot of the tree among the moss-covered roots of which it has passed the winter in the pupal state. The female moth seldom lives for more than three or four days; but the males which fly swiftly in the forests, seeking for the less abundant females, live for a much longer period, certainly from eight to fourteen days[2 - This estimate is derived from observation of the time during which these insects are to be seen upon the wing. Direct observations upon the duration of life in this species are unknown to me.].

The females of the Psychidae also deposit all their eggs in one place. The grasses and lichens upon which their caterpillars live grow close at hand upon the surface of the earth and stones, and hence the female moth does not leave the ground, and generally does not even quit the pupa-case, within which it lays its eggs; as soon as this duty is finished, it dies. In relation to these habits the wings and mouth of the female are rudimentary, while the male possesses perfectly developed wings.

The causes which have regulated the length of life in these cases are obvious enough, yet still more striking illustrations are to be found among insects which live in colonies.

The duration of life varies with the sex in bees, wasps, ants, and termites: the females have a long life, the males a short one; and there can be no doubt that the explanation of this fact is to be found in adaptation to external conditions of life.

The queen-bee—the only perfect female in the hive—lives two to three years, and often as long as five years, while the male bees or drones only live four to five months. Sir John Lubbock has succeeded in keeping female and working ants alive for seven years—a great age for insects[3 - [Sir John Lubbock has now kept a queen ant alive for nearly 15 years. See note 2 {note 18 below} on p. 51.—E. B. P.]],—while the males only lived a few weeks.

These last examples become readily intelligible when we remember that the males neither collect food nor help in building the hive. Their value to the colony ceases with the nuptial flight, and from the point of view of utility it is easy to understand why their lives should be so short [See Note 7 (#x9_pgepubid00014) and Note 9 (#x9_pgepubid00016)]. But the case is very different with the female. The longest period of reproduction possible, when accompanied by very great fertility, is, as a rule, advantageous for the maintenance of the species. It cannot however be attained in most insects, for the capability of living long would be injurious if all individuals fell a prey to their enemies before they had completed the full period of life. Here it is otherwise: when the queen-bee returns from her nuptial flight, she remains within the hive until her death, and never leaves it. There she is almost completely secure from enemies and from dangers of all kinds; thousands of workers armed with stings protect, feed, and warm her; and in short there is every chance of her living through the full period of a life of normal length. And the case is entirely similar with the female ant. In neither of these insects is there any reason why the advantages which follow from a lengthened period of reproductive activity should be abandoned [See Note 6 (#x9_pgepubid00013)].

That an increase in the length of life has actually taken place in such cases seems to be indicated by the fact that both sexes of the saw-flies—the probable ancestors of bees and ants—have but a short life. On the other hand, the may-flies afford an undoubted instance of the shortening of life. Only in certain species is life as short as I have indicated above; in the majority it lasts for one or more days. The extreme cases, with a life of only a few hours, form the end of a line of development tending in the direction of a shortened life. This is made clear by the fact that one of these may-flies (Palingenia) does not even leave its pupa-skin, but reproduces in the so-called sub-imago stage.

It is therefore obvious that the duration of life is extremely variable, and not only depends upon physiological considerations, but also upon the external conditions of life. With every change in the structure of a species, and with the acquisition of new habits, the length of its life may, and in most cases must, be altered.

In answering the question as to the means by which the lengthening or shortening of life is brought about, our first appeal must be to the process of natural selection. Duration of life, like every other characteristic of an organism, is subject to individual fluctuations. From our experience with the human species we know that long life is hereditary. As soon as the long-lived individuals in a species obtain some advantage in the struggle for existence, they will gradually become dominant, and those with the shortest lives will be exterminated.

So far everything is quite simple; but hitherto we have only considered the external mechanism, and we must now further inquire as to the concomitant internal means by which such processes are rendered possible.

This brings us face to face with one of the most difficult problems in the whole range of physiology,—the question of the origin of death. As soon as we thoroughly understand the circumstances upon which normal death depends in general, we shall be able to make a further inquiry as to the circumstances which influence its earlier or later appearance, as well as to any functional changes in the organism which may produce such a result.

The changes in the organism which result in normal death,—senility so-called,—have been most accurately studied among men. We know that with advancing age certain alterations take place in the tissues, by which their functional activity is diminished; that these changes gradually increase, and finally either lead to direct or so-called normal death, or produce indirect death by rendering the organism incapable of resisting injuries due to external influences. These senile changes have been so well described from the time of Burdach and Bichat to that of Kussmaul, and are so well known, that I need not enter into further details here.

In answer to an inquiry as to the causes which induce these changes in the tissues, I can only suggest that the cells which form the vital constituents of tissues are worn out by prolonged use and activity. It is conceivable that the cells might be thus worn out in two ways; either the cells of a tissue remain the same throughout life, or else they are being continually replaced by younger generations of cells, which are themselves cast off in their turn.

In the present state of our knowledge the former alternative can hardly be maintained. Millions of blood corpuscles are continually dying and being replaced by new ones. On both the internal and external surfaces of the body countless epithelial cells are being incessantly removed, while new ones arise in their place; the activity of many and probably of all glands is accompanied by a change in their cells, for their secretions consist partly of detached and partly of dissolved cells; it is stated that even the cells of bone, connective tissue, and muscle undergo the same changes, and nervous tissue alone remains, in which it is doubtful whether such a renewal of cells takes place. And yet as regards even this tissue, certain facts are known which indicate a normal, though probably a slow renewal of the histological elements. I believe that one might reasonably defend the statement,—in fact, it has already found advocates,—that the vital processes of the higher (i.e. multicellular) animals are accompanied by a renewal of the morphological elements in most tissues.

This statement leads us to seek the origin of death, not in the waste of single cells, but in the limitation of their powers of reproduction. Death takes place because a worn-out tissue cannot for ever renew itself, and because a capacity for increase by means of cell-division is not everlasting, but finite [See Note 8 (#x9_pgepubid00015)]. This does not however imply that the immediate cause of death lies in the imperfect renewal of cells, for death would in all cases occur long before the reproductive power of the cells had been completely exhausted. Functional disturbances will appear as soon as the rate at which the worn-out cells are renewed becomes slow and insufficient.

But it must not be forgotten that death is not always preceded by senility, or a period of old age. For instance, in many of the lower animals death immediately follows the most important deed of the organism, viz. reproduction. Many Lepidoptera, all may-flies, and many other insects die of exhaustion immediately after depositing their eggs. Men have been known to die from the shock of a strong passion. Sulla is said to have died as the result of rage, whilst Leo X succumbed to an excess of joy. Here the psychical shock caused too intense an excitement of the nervous system. In the same manner the exercise of intense effort may also produce a similarly fatal excitement in the above-mentioned insects. At any rate it is certain that when, for some reason, this effort is not made, the insect lives for a somewhat longer period.