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syahdu-brodin OM BARBARA Ft ki ageng slamet There is no doubt that Einreiseformalitäten Usa reproduce much more Victorious during the embryonic period. The purely physiological conditions which determine this limit leave room for a considerable amount of variation in longevity. As a result of this displacement of lime in old age, the blood-vessels become modified in a distinctive fashion. Posts to:. Wiesbaden,pp. Only the caecum can be thought to have some digestive property.

The same observer, it is true, has shown that the hair of young men of between 21 and 24 years grew at the rate of 15 mm.

Pohl himself has shown that, in the latter region, the hair grows slower than in other regions. Moreover, in many boys of 11 to 15 years old, studied by this observer, the rate of growth was always less than 15 mm.

I have been able to note that the nails grow even in very old people. In the case of Mme. In the case of a lady of 32 years old, the corresponding nail grew 3 mm.

The centenarian's nails had to be cut from time to time. Although the hairs of old people grow, they become white, which is a phenomenon of senile degeneration.

Although they increase in length, the colouring matter in them becomes reduced and finally disappears.

In the " Nature of Man " I described the process by which this blanching takes place, and which may now be regarded as definitely proved.

It is useful as a means of interpreting the real nature of the process of senescence. In several published works, I have explained my belief that just as the pigment of the hair is destroyed by phagocytes, so also the atrophy of other organs of the body, in old age, is very frequently due to the action of devouring cells which I have called macrophags.

These are the phagocytes that destroy the higher elements of the body, such as the nervous and muscular cells, and the cells of the liver and kidneys.

This part of my theory has encountered very strong criti- cism, especially with regard to the part played by the macrophags in the senescence of nervous tissue.

For several years M. Marinesco 1 has attacked my theory of the atrophy of the nerve-cells in old age. In the first place, he has stated that in old people, and even if these are very old, it is rare to find phagocytes surrounding and devouring the cells of the brain.

In support of this contention, he has been good enough to send me two pre- parations made from the brains of two very old persons. After careful examination I was convinced that my oppo- nent had been inexact.

In the brain of the two centenarians one of whom died at the age of 1 17 years there were very many nerve-cells surrounded by phagocytes and in process of being destroyed by them.

It happened, however, that as the sections were very weakly stained, it was more diffi- cult to observe the facts than in the preparations upon which I had made my own observations.

I have already recorded this fact in the second and third French editions of the " Nature of Man. Marinesco has published another criticism of my theory in an article' 2 entitled " Histological Investigations into the Mechanism of Senility.

He thinks that nerve-cells atrophy independently of the cells that surround them. The latter, the so-called neur- onophags, only contribute to the atrophy inasmuch as they press against the nerve-cells and deprive them of nutrition.

He is confident that the constituent parts of nerve-cells are never found in the neuronophags. Leri has taken a similar view in a Report on the Senile Brain 1 presented to a recent congress of alienists and neurologists.

According to him " the nuclei which surround some of the atrophying nerve-cells do not play the part of neuronophags. Sand elaborates the same view. He relies on his observation that " neuronophags are usually either devoid of protoplasm or display only a very thin layer of it.

They never exhibit protoplasmic outgrowths, and they never have granules in their cellular bodies p. Laignel-Lavastine and Voisin a have taken the same view, maintaining that the neurono- phags do not display phagocytosis.

Although I cannot undertake here to give a detailed reply to the arguments of my critics, I may point out a fallacy that vitiates their reasoning.

The study of the intimate structure of nervous tissue involves the treatment of that very delicate substance by numerous active reagents. It is extremely important not to forget the possibility of altera- tions which may be produced in the processes of preparation and which are extremely difficult to avoid.

A glance at the figures given by my critics shows me that the neurono- phags in their preparations had been subjected to violent treatment. When M.

Leri speaks of "the nuclei which surround some of the nerve-cells," and M. Sand of " cells without protoplasm," it is clear that they had been observ- ing cells destroyed by the processes of the laboratory.

The 1 Le Bulletin mtdical, , p. Marinesco show that in his preparations, too, the neuronophags had been very greatly altered.

It is well known that nuclei do not exist free in tissues, and that when they appear devoid of protoplasm, there has been some defect in the technical methods of preparing them for examination.

As a matter of fact, neuronophags do not consist of nuclei with at the most a pellicle of proto- plasm ; like other cells, they have protoplasmic bodies which, however, are frequently destroyed by the violent processes of histological preparation.

The arguments of my critics recall to me the words of a medical student, who, on being asked to describe the microbe of tuberculosis, said that it was a little red bacillus.

The bacillus in question, like most bacilli, is colourless, but it is usual to stain it so that it may be visible under the microscope.

The student, knowing it only in particu- lar preparations, had a false idea of its appearance. In well-made preparations, neuronophags are typical cells with abundant protoplasm.

When they have been pre- served by a process that does not dissolve their contents, they show granules like those found in nerve-cells.

To study neuronophagy, M. He succeeded in showing first that in the destruction of nerve-cells that occurs in cases of hydrophobia, the contents of these cells are absorbed by the surrounding neuronophags.

What becomes of these granulations on the destruction and dis- appearance of the nerve-cell? If, as M. Marinesco has it, there is no phagocytosis by the surrounding cells, but merely a mechanical interference, then the granules, on the destruction of the nerve-cells that contained them, should be found lying in the interstitial tissue.

But this does not happen. The granules are ingested by cells which are true macrophags. Manouelian has shown that in the case of senile brains the granules of the nerve-cells are absorbed by neurono- phags.

I have myself studied M. Manouelian's prepara- tions and can testify to the accuracy of his observations Figs. Doubt is no longer possible.

In senile degeneration the nerve-cells are surrounded by neuronophags which absorb their contents and bring about more or less complete atrophy.

It has been supposed that in order to devour their contents, the neuronophags must penetrate the nerve-cells, and such an event has rarely been seen.

But it is well known, the phagocytosis of red blood corpuscles being a typical instance, that to absorb a cell a phagocyte does not necessarily engulf it bodily or penetrate it, but may gradu- ally denude it of its contents merely by resting in contact with it.

There has been some discussion as to the condition of nerve-cells which are on the point of being devoured by neuronophags.

It has been noticed that such cells may display a considerable amount of degeneration without being devoured, whilst, on the other hand, cells apparently normal have been found undergoing phagocytosis.

Although the destruction of nerve-cells by neuronophags is a general occurrence in senile brains, one may conceive of cases where this does not occur.

And so, in old people who have preserved their faculties, it may well be that the neuronophags have refrained from attacking the nerve-cells.

Two nerve-cells from the cortex of the brain of an old dog aged fifteen years. The neuronophags surrounding the nerve-cells contain numerous granulations.

From preparations made by M. But as such instances are rare, so also phagocytosis is usually found in senile brains, and I cannot accept M.

Sand's denial of its existence, based on his study of two cases. Weinberg have completely supported this view.

The bleaching of hair and the atrophy of the brain in old age thus furnish important arguments against the view that senescence is the result of arrest of the reproduc- tive powers of cells.

Hairs grow old and become white without ceasing to grow. The cessation of the power of reproduction cannot be the cause of the senescence of brain- cells, for these cells do not reproduce even in youth.

The blanching of hair is due to the destructive agency of chromophags; in atrophy of the brain neur- onophags destroy the higher nerve-cells.

In addition to these instances of phagocytosis, in which the active agents belong to the category of macrophags, there are many other devouring cells, adrift in the tissues of the aged, and ready to cause destruction of other cells of the higher type.

The phagocytic action is not so manifest as in the case of infec- tious diseases, partly because it is the method of macro- phags to absorb the contents of the higher cells extremely slowly.

The mode of action is well seen in the atrophy of an egg-cell Fig. Matchinsky l has studied the series of events in my laboratory, and I am myself well assured of the importance of the action of macrophags in the atrophy of the ovary.

The phenomena of atrophy in general and of senile decay afford other cases of tissue destruction in which the phago- FIG. Ovum of a Bitch in process of destruction by Phagocytes, which are full of fatty granules.

After M. It is well known that progressive muscular debility is an accompaniment of old age. Physical work is seldom given to men over sixty years of age, as it is notorious that they are less capable of it.

Their muscular movements are feebler and soon bring on fatigue ; their actions are slow and painful. Even old men whose mental vigour is un- impaired admit their muscular weakness.

The physical 1 Annales de FInstitut Pasteur, , vol. More than half a century ago, Kolliker, 1 one of the founders of histology, devoted some attention to this matter, and described the senile modification of muscular tissue in the following words : " In old age there is a true atrophy of the muscles.

The fibres are much more slender ; there are deposited in their substance numerous yellow or brown granules and many globular nuclei.

These nuclei are fre- quently arranged in longitudinal series and present such signs of active division as are found in embryonic tissue. Vulpian 3 and Douaud 3 have stated that a multiplication of nuclei takes places in the atrophying muscles of the old.

As the senile degeneration of muscular tissue appeared to be important in my study of the mechanism of senescence, M. Weinberg and I examined several qtases of muscular atrophy in old human beings and lower animals.

We were able to recognise the phenomena observed by our prede- cessors. In senile atrophy the muscular fibres contain many nuclei, and these, increasing rapidly, bring about an almost complete disappearance of the contractile substance Fig.

The fibres preserve their striation for a certain time but eventually lose it and appear to contain an amor- phous mass with numerous, rapidly multiplying nuclei.

The investigators who had recorded these facts thought of them only as curious. It is plain, in the first place, however, that this remarkable and rapid multiplication is a proof that senile atrophy is not due to failure of cell pro- 1 Elements fhistologie humaine, French translation, , p.

Paris, In muscular atrophy, cell- multiplication, so far from failing, greatly increases. We may add muscular atrophy to the blanching of hair and the decay of nerve-cells as another instance showing that senile degeneration is not the result of cells ceasing to be able to FIG.

Degeneration of striated muscle Fibres from the auricular muscle of a man aged 87 years. From a preparation made by Dr. Just as in the atrophy of the brain there is an increase in the volume of neurogloea, the substance in which the neuronophags are found, so also in the atrophy of the muscles there is an increase of muscular nuclei.

Along with the increase of nuclei, however, there is an increase of the protoplasmic substance of the fibres known as sarco- plasm. In a normal muscle the two substances and the sarcoplasmic nuclei are in equilibrium, but in old age the sarcoplasm and its nuclei increase at the expense of the myoplasm.

The equilibrium is destroyed with the result that the muscular power is weakened. In these conditions the sarcoplasm acts phago- cytically with regard to the myoplasm, just as the chromo- phag becomes the phagocyte of the pigment of the hair, or the neuronophag devours the nerve-cell.

The investigation of other cases of muscular atrophy, as, for instance, that of the caudal muscles of frog-tadpoles, confirms the significance of the process that I have observed in old age.

In the two cases, what takes place is the destruction of the contractile material of the muscles by myophags, a special kind of phagocyte.

It is one of the curiosities of senile atrophy that whilst there is hardening or sclerosis of so many organs, the skele- ton, the most solid part of our frame-work, becomes less dense, so that the bones are friable, the condition often leading to serious accidents in old people.

The bones become porous, and lose weight. It is difficult to believe that macrophags, although they destroy softer elements such as nerve-cells or muscle fibres, can be able to gnaw through a hard material like bone impregnated with mineral salts.

As a matter of fact, the mechanism of bone atrophy must be placed in a different category from the phagocytosis of other organs. It is brought about, how- ever, by the agency of cells very like some of the macro- phags.

These cells contain many nuclei, and are known as osteoclasts. They form round about the bony lamellae and lead to their destruction, but are incapable of breaking off fragments of bone and dissolving them in their interiors.

The process can be observed in the different varieties of caries of the bone, and in the bony atrophy of old age as is represented in Fig.

By the action of the osteoclasts, which themselves are macrophags, part of the lime in the skeleton is dissolved during old age and passes into the general circulation.

This is probably a source of the lime which is deposited so readily in the different tissues of old people.

Whilst the bones become lighter, the cartilages become bony, the inter- FlG. Destruction by osteoclasts of bony matter in the sternum of a man aged 8 1 years.

As a result of this displacement of lime in old age, the blood-vessels become modified in a distinctive fashion. Atheroma of the arteries is not invariable in old people, but it occurs extremely frequently.

In this form of degenera- tion, lime salts are deposited in the walls of the cells, so that they become hard and friable. Several others, among whom I may mention Durand-Fardel and Sauvage, have laid stress on the coincidence of atheromatous lesions of the arteries and senile degeneration of the bones.

The relations. It is the manifestation of an extraordinary disturbance of the properties of the cells that compose the body.

The atheromatous condition of the arteries is closely linked with arterial sclerosis, an affection which is very common, although not constant, in the aged.

The whole question of these vascular alterations is extremely complex, and before it can be cleared up, a number of special inves- tigations must be made.

Probably diseases of the arteries of different kinds, and arising from different causes, are grouped under the terms atheroma and sclerosis. In some cases the lesions are in- flammatory and are due to the poisons of microbes.

An example of such an origin is the case of syphilitic sclerosis, in which the specific microbes spirilla of Schaudinn lead to precocious senescence.

In other cases the arteries show phenomena of degeneration resulting in the formation of calcareous platelets which interfere with the circulation of the blood.

Investigations which have been made in recent years have led to very interesting results concerning the origin of atheroma of the arteries.

In most cases, attempts to produce such lesions of the arteries by experimental 1 Demange, tude sur la vieillesse, , p.

Josue" 1 has been able to produce true arterial atheroma in rabbits by injecting into them adrenaline, the secretion of the suprarenal cap- sules.

This experiment has been repeated many times and is now well known. Later on, M. Boveri 2 obtained a similar result by injecting nicotine, the poison of tobacco.

It is obvious, therefore, that amongst the arterial diseases which play so great a part in senescence, some are chronic inflam- mations produced by microbes, whilst others are brought about by poisons introduced from without.

It is easy to understand, therefore, why these diseases of the arteries are not always present in old age, although they are very common. The part played by the secretion of the suprarenal glands in the production of arterial disease has brought renewed attention to a theory which supposed that certain glandular organs in the body play a preponderating part in senile degeneration.

Lorand 3 in particular has argued that " senility is a morbid process due to the degeneration of the thyroid gland and of other ductless glands which normally regulate the nutrition of the body.

Everyone who has seen the cretins in Savoy, Switzerland, or the Tyrol, must have noticed the aged appearance of these victims, although very often they are quite young.

The condition of cretinism, with its pro- found bodily changes, is the result of degeneration of the thyroid gland.

On the other hand, it is well known that 1 C. It is quite probable, therefore, that these so-called vascular glands have their share in producing senility.

Many facts show that they destroy certain poisons which have entered the body, and it is easy to see that, if they have become functionless, the tissues are threatened with poisoning.

It does not follow, how- ever, that their action in producing senility is exclusive, or even preponderating.

Weinberg, at the Pasteur Insti- tute, made special investigations on this point, and found that the thyroid gland and the suprarenal capsules were almost invariably normal in old animals cat, dog, horse , although the latter showed unmistakable signs of senility.

Similarly in an old man of 80 years, who died from pneu- monia, the thyroid gland was quite normal. It must not be forgotten that the aged very often die from infectious diseases such as pneumonia, tuberculosis, and erysipelas.

In these diseases the vascular glands generally, and the thyroid gland in particular, are very often affected, with the. In the masterly accounts of such unfortunates, recently compiled by the well-known surgeon Kocher 2 there are many points which are char- acteristic, without being typical, of old people.

Oedema of the skin which characterises thyroid patients 1 Sarbach, Mittheilungen a. Wiesbaden, , pp.

The loss of hair, normal in the patients, is not a character of old age. In myx- edematous women, menstruation is very active; it ceases in old women.

The great muscular development of myxedematous patients distinguishes them from old people. Physiological investigation does not support the exist- ence of any strong affinity between old age and affection of the thyroid gland.

Bourneville and Bricon l having shown that the tendency to cachexia after extirpation of the thyroid ceases almost abruptly at the age of thirty.

That age may be taken as the limit of youth, of the time when growth is vigorous and the function of the thyroid most active.

Cases of cachexia, where the thyroid gland has been removed in old persons from fifty to seventy, are very. Rodents rats, rabbits support the removal of the thyroid extremely well, without signs of cachexia, although these are normally short-lived creatures.

According to Horsley 2 extirpation of the thyroid is not followed by cachexia in birds or rodents and is followed by it only very slowly in ruminants and horses ; it produces the condition invariably but slightly in man and monkeys and extremely seriously in carnivora.

If this series be compared with the informa- tion given in the next section of this volume on the relative ages which the animals in question attain, it will be seen that there is no correspondence.

In short, whilst I do not deny that the vascular glands may take a share in the causation of senility, in so far as 1 Archives de Neurologie, I think it indubitable that in senescence the most active factor is some alteration in the higher cells of the body, accompanied by a destruction of these by macrophags which gradually usurp the places of the higher elements and replace them by fibrous tissue.

Such a process affects the organs of secretion kidneys , the reproductive organs, and in a modified form the skin, the mucous membranes, and the skeleton.

The testes are amongst the organs which resist invasion by macrophags. I have already given an ex- ample " The Nature of Man," p. I know of a similar case, the age being years.

Such cases are not rare, and not only in old men, but in old animals, the testes continue to FIG. Testis tissue from a dog aged twenty-two years.

Wemberg and From a preparation made by Dr. I have investigated these Weinberg. Many of the organs of the animal exhibited serious invasions by macrophags but the testes were extremely active, the cells being in free proliferation and producing abundant spermatozoa Fig.

In harmony with this condition of the sexual organs, the sexual instincts of the animal remained normal. We have investigated another dog which died at the age of eighteen years.

In this case the testes were cancerous and there was no possibility of the production of spermatozoa. It is manifest that the tissues do not invariably degene- rate in old age, nor do all the organs that are modified in old age show destruction by phagocytes and replacement by connective tissue.

Organs which produce phagocytes, such as the spleen, the spinal marrow and the lymphatic glands, certainly show traces in old age of fibrous degenera- FIG.

An old dog, iged eighteen years. I have frequently noticed cell division in such organs, and as an example inay give the case of the bone marrow taken from a man of 81 years Fig.

The eye is an organ that is modified in old age without the action of macrophags. These modifications are due to impregnation of the parts affected by fatty matter which makes them opaque.

This deposition of fat 1 has been attributed to defective nutrition. In most organs such fatty degeneration is followed by phagocytosis, but the cornea and the crystalline lens are exempt from this conse- quence for anatomical reasons.

Most organs possess in addition to their higher elements a constant source of macrophags. Such a source of phagocytosis is the neuro- glcea in nervous tissues, the A sarcoplasm in muscular tis- sues ; the bones contain osteo- clasts and the liver and the kidneys are readily invaded by phagocytes from the blood.

The lens and the cornea have no cells that are able to become macrophags. A syphilitic child is "a Weinberg. It is no mere analogy to suppose that human senescence is the result of a slow but chronic poisoning of the organism.

Such poisons, if not completely destroyed or eliminated, weaken the tissues, the functions of which become altered or enfeebled, so that, 1 Fuss, Der Greisenbogen, in Virchotv's Archi-v, , vol.

Toufesco, Sur le cristallin, Paris, The phagocytes resist the influence of invading poisons better than any of the other cells of the body and some- times are stimulated by them.

The general result of such conditions is that there comes to be a struggle between the higher cells and the phagocytes in which the latter have the advantage.

The answer to the question as to whether our senescence can be ameliorated must be approached from several points of view. This course I shall now follow.

Some, as for instance, the males of certain wheel animalculae Rotifera complete their cycle of life from birth to death in 50 or 60 hours, whilst others, like some reptiles, live more than 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.

Buffon 1 long ago stated his opinion that the " total duration of life bore some definite relation to the length of the period of growth.

Just as any species has acquired a fixed and practically invariable size, so it would have acquired a definite longe- vity. 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 dura- tion 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 roo 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.

Flourens 2 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 them- 1 Histoire natitrelle gtntrale et particultire, vol.

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, years ; the camel takes 8 to grow, and lives 5 times as long, i.

However, even if we consider only the mammalia, it is impossible to accept Flourens' law, without considerable reserve.

Weismann l 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 reach- ing 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.

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 repro- duce, 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 1 Ueber die Dauer des Lebens, Jena, , p. 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 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.

Milne-Edwards 1 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.

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.

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. 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 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.

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 circum- stances the duration of life becomes a factor in the preserva- tion 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 Jead 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 years, or live beyond that time, much more often than men.

There are parrots which lay two or three times a year, producing six to nine eggs in each clutch. The ducks Anatidae 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 carni- vora. The longevity is possibly to be explained as an adaptation for the preservation of the species by compen- sating 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 in- dependently.

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.

Oustalet, 1 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.

Elephants and parrots, for instance, are vegetarian and reach very great ages. On the other hand, there exist long-living carni- vorous 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.

As the higher animals are nearly always larger than invertebrates, if there be a definite relation between long- evity and size, one would expect to find that vertebrates live longer than invertebrates.

However, this is not the case. Amongst animals of extremely simple organisation, there are some which reach a great age. A striking example of this is found in sea-anemones.

These animals have a very simple structure, without a separate digestive canal, and with a badly developed, diffused nervous system, and yet have lived very long in captivity.

More than forty years ago, I remember having seen in the possession of M. Lloyd, the Director of the Aquarium at Hamburg, an anemone that he had kept alive for several dozen years 'in a glass bowl.

Another sea-anemone, belonging to the species Actinia mesembryanthemum, is known to have lived 66 years. It survived its owner for 36 years, and died in Edinburgh in , the cause of death being un- known.

Although they are thus capable of living so long, the rate of growth of members of this species is rapid, and their fertility is very high. According to Dalyell, these anemones reach the adult condition in 15 months.

The specimen in his possession, in the 20 years from to produced larvae, then after a period of sterility it gave birth, in one night to 2 3 young anemones.

This extraordinary prolificness decreased with age, but even when it was 58 years old it used to produce from 5 to 20 at a time. In the seven years from onwards, it gave birth to young anemones.

Ashworth and Nelson Annandale have published their observations on another sea-anemone, of the species Sagartia troglodytes, which was 50 years old.

It differed from younger examples only in being less prolific. There are other polyps, such as Flabellum, which do not live more than 24 years, although we have no knowledge as to the cause of the different duration of life.

The variation in the length of the life of molluscs and insects is extremely great. Some species of gasteropods Vitrina, Succinea live only a very few years, whilst others Natica heros can reach thirty years.

Some of the marine bivalves, as for instance, Tridacna gigas, can live to sixty or a hundred years. Some live only a few weeks; 1 Ashworth and Annandale, Proceedings of the R.

Society of Edinburgh, vol. Ordnungen des Thierreichs, vol. In the same order of Insects, however, Hemiptera there are species of cicada which live thirteen to seventeen years, that is to say, much longer than such little Rodents as rats, mice, and guinea-pigs.

The larva of an American species spends seventeen years buried in the ground in orchards, where it feeds on the roots of apple trees, and the species is known as Cicada septemdecim, because of this duration of life.

In the adult stage the insect lives little more than a month, just time enough to lay the eggs, and bring into the world the new generation, which in its turn will not appear above ground until after another period of seven- teen years.

Between these extremes of long and short life, there is to be found amongst insects almost every gradation of longevity.

Science, in its present state, has failed to find any law governing these facts. Rules which hold good up to a certain point in the case of the higher animals break down in their application to insects.

The large grass- hoppers and locusts, for instance, live a much shorter time than many minute beetles. Queen bees, the fertility of which is very great, live two or three years and may reach a fifth year, whilst worker bees, which are infertile, die in the first year of their existence.

Female ants, although these are small and extremely prolific, reach the age of seven years. It is more probable that we shall find some explanation in the case of vertebrates concerning which we know much more.

As a general rule, it may be laid down that the lower vertebrates live longer than mammals. The facts about the longevity of fish are not very numer- ous, but it seems clear that these animals reach a great age.

The ancient Romans, who used to keep eels in aquaria, have noted that these fish would live for more than sixty years.

There is reason to believe that salmon can live for a century, whilst pike live much longer. There is, for instance, the much quoted instance of the pike stated by Gessner to have been captured in and to have lived for years afterwards.

Carps are regarded as equally long lived, Buffon setting down their period of life as years. There is a popular idea that the carp in the lakes at Fontainebleau and Chantilly are several centuries old, but E.

Blanchard throws doubt on the accuracy of this esti- mate, inasmuch as during revolutionary times most of the carp were eaten when the palaces were overrun by the popu- lace.

There is no doubt, however, that the life of carp may be very long indeed. Not very much is known about the duration of life in batrachians, but it is certain at least that some small frogs may live twelve or sixteen years, and toads as many as thirty-six years.

More is known about the life of reptiles. Crocodiles and caymans, which are large and which grow very slowly, attain great ages. In the Paris Museum of Natural History there are crocodiles which have been kept for more than forty years without showing signs of senescence.

Turtles, although they are smaller than crocodiles, live still longer. Another tortoise, a native of the Galapagos Islands, is known to be years old, whilst a specimen in the London Zoological Gardens is years old.

A land tortoise Testudo marginata has been kept in Norfolk, England, for a century. I am informed that in the Archbishop's palace at Canterbury, there is to be seen the carapace of a tortoise which was brought to the Palace in and which lived there for years.

I have already referred to a specimen of Testudo mauritanica, the history of which is known for 86 years, but which is probably much older.

Very little is known as to the longevity of lizards and serpents, but it may be inferred from what I have said about other reptiles that reptiles as a class are able to reach great ages.

It is an easy inference that the great duration of life in cold-blooded animals is associated with the slowness of the physiological processes in these creatures.

The circu- lation, for instance, is so slow, that the heart of a tortoise beats only 20 to 25 times in a minute.

Weismann has suggested that one of the factors influencing the duration of life is the rapidity or slowness of the vital activities, the times taken by the processes of absorption and nutrition.

On the other hand, the blood is hot and the vital activities are rapid in birds, and yet birds may attain great ages. Although in the last chapter I gave a number of examples, the subject is so important that I propose to go further into details.

The possibility of this is due to an admirable set of details brought together by Mr. Canaries have lived in cap- tivity from 17 to 20 years, and goldfinches up to 23 years.

Field larks have lived for 24 years, the Lesser Black-backed Gull 31 years and the Herring Gull 44 years. Birds of medium size may live for several dozens of years, whether they live on animal or on vegetable food, whether they are prolific or lay very few eggs.

I will quote only a few instances. Of forty parrots the minimum and maximum ages were respectively 15 and 81 years, and the average 43 years.

Without accepting the truth of the story mentioned by Humboldt according to which certain parrots survived an extinct race of Indians, at least we may be certain that great ages have sometimes been reached by these birds.

Levaillant mentions a parrot Psittacus erithaceus which lost its memory at the age of 60 years, its sight at 90 years, and which died aged 93 years.

Another individual, probably of the same species, is reported by J. Jennings to have reached the age of Jones, Layard, and Butler are the authorities for instances of Sulphur-crested Cocka- toos having reached respectively 30, 72 and 81 years.

Abrahams states that an Amazon Chrysotis amasonica lived 1 02 years. I myself have observed two cases of great longevity in the same species of parrot.

One of these birds died at the age of 82 years, apparently simply from old age, whilst the other, which was in my possession for several years before it died at the age of 70 to 75 years, was vigorous, showing no signs of senility, but died of pneu- monia.

One raven reached 69 years and another 50, an Eagle-owl Bubo maximus 68 years, another 53, a condor 52, an imperial eagle 56, a com- mon heron 60, a wild goose 80, and a common swan 70 years.

None of these examples approaches the legendary three centuries attributed to the swan, but it is evident that many different kinds of birds may attain great age.

I can add some cases to those of Mr. In the Royal Park at Schonbrunn, near Vienna, a white-headed vulture Neo- phron percnopterus died aged years, a golden eagle Aquila chrysaetus aged , and another aged 80 accord- ing to Oustalet.

Pycraft Country Life, June 25th, reported that a female eagle, captured in Norway in , had been brought to England and had lived for 75 years.

In the last thirty years of its life, it had produced ninety eggs. The same writer mentions the case of a falcon having lived to years.

The collection of facts that I have passed in review make it manifest that birds may have a great duration of life, but that reptiles surpass them in this respect.

Birds cer- tainly do not reach the very great ages of crocodiles and tortoises. Longevity, therefore, is reduced as we ascend in the scale of vertebrate life.

We find a still greater reduction when we turn from birds to mammals. Some mammals, it is true, may live as long as birds. Elephants are a good instance.

It used to be thought that these giant mammals could live three or four centuries, but I can find no confirmation of the legend, which seems as mythical as that relating to the life of swans.

There are no exact data as to the ages reached by wild elephants, but it has been stated that in captivity an elephant rarely but occa- sionally has completed its century.

In the official list of the Indian Government, which gives the fteaths of elephants, it appears that of examples, only one lived more than 20 years after it had been purchased Brehm's Mammals.

Flourens, using his own formula, assigned the age of years to elephants as their epiphyses do not fuse with the long bones until the age of So far, I know 7 of no fact to support the conclusion, although it seems fairly well established that occasionally an elephant may reach a century.

It is stated that one elephant was in service throughout the whole period of more than years in which Ceylon was occupied by the Dutch.

This elephant was found in the stables in Natives with special knowledge of elephants set down their duration of life as from 80 to years, but say that they begin to grow old at from 50 to 60 years of age.

My general conclusion from the facts is that the life of these very large mammals is about the same as that of man who is very much smaller.

Centenarians, extremely rare amongst elephants, do not appear to exist in any other kind of mammals except man.

The rhinoceros, another large mammal which is a native of the same countries as the elephant, does not reach a great age. According to Oustalet an Indian rhinoceros died in the menagerie of the Paris Museum at about the age of 25 years, and showed all the signs of senility.

Another Indian rhinoceros lived for 37 years in the London Zoological Gardens. Horses and cattle are large animals, hut do not enjoy very long lives.

The usual duration of life in horses is from 15 to 30 years. They begin to grow old about 10 years, and in very rare cases may reach 40 or more.

A Welsh pony is said to have reached the age of sixty, but such a case is excessively rare. Two other extreme cases are that of a horse belonging to the Bishop of Metz which died at the age of 50 years, and the charger of Field- Marshal Lacy which died at The duration of life of cattle is still shorter.

Domestic cattle show the first sign of age, a yellow discoloration of the teeth, when five years old. In the sixteenth to eighteenth year the teeth fall out, or break, and the cow ceases to give milk, whilst the bull has lost reproductive power.

According to Brehm, cattle live for 25 to 30 years or more. Although the duration of life is short, cattle are not prolific.

The gestation period of a cow approaches that of the human race days , and there is only one birth a year. The total period of reproductivity lasts only a few years.

The sheep, another domesticated Ruminant, has a life even shorter. According to Grindon, sheep do not live longer than 12 years as a rule, but may reach 14 years, which in their case would be extreme age, as they generally lose their teeth at from 8 to 10 years.

Some Ruminants, such as camels and deer, apparently live longer than sheep or cattle, but I do not know exact facts about them.

The short life of domesticated carnivorous animals is well known. Jonatt has mentioned as an extreme rarity a dog of 22 years of age, and Sir E. Ray Lankester Comparative Longevity, p.

The oldest dog that I have been able to procure died at the age of It is generally believed that cats do not live so long as dogs.

The average age which they may attain is usually thought to be 10 or 12 years, but certainly a cat of that age has not the decrepid appearance of an old dog.

Thanks to the kindness of M. Barrier, the Director of the Ecole d'Alfort, I have had in my possession a cat 23 years old.

It appeared to be quite vigorous, and died from cancer in the liver. Most rodents, particularly the domesticated kinds, are extremely prolific and very short lived.

It is extremely rare for a rabbit to reach the age of 10 years, whilst 7 years is the utmost limit for a guinea-pig. Mice, so far as I can ascertain, do not live more than 5 or 6 years.

It is plain from the facts that I have brought together, that mammals, whether they are large or small, as a rule, have shorter lives than birds.

It is probable, therefore, that there is something in the structure of mammals which has brought about a shortening in the duration of their lives.

Whilst most of the lower vertebrates, and all birds, reproduce by laying eggs, the vast majority of mammals are viviparous.

As the tax on the parent organism is greater when the young are produced alive than when eggs are laid, it might be thought that in this difference lay the cause of the shorter life of mammals.

There are many facts, however, which make it impos- sible to accept such a view. The longevity of mammals is nearly equal in the two sexes, although the tax on the organism caused by reproduction is much greater in the case of females than in males.

Longevity, however, cannot be regarded as a character stable in each species and necessarily identical in the two sexes. The animal kingdom presents many cases of disparity in this respect, the difference in longevity in the tw ; o sexes being specially striking in species of insects.

Generally, the females live longer than the males, as, for instance, amongst the Strep- si ptera, where the females have 64 times the duration of life of the males.

On the other hand, amongst butterflies, there are cases e. In the human race, there is a difference in the longevity of the sexes, the females having the advantage.

As in most cases of disparity in the duration of life the female lives longer than the male, it is plain that the differ- ence cannot be assigned to the drain on the organism caused by reproduction, which, of course, is much greater in females.

Moreover, a closer scrutiny of the facts shows that although mammals do not live so long as birds, the repro- ductive drain is greater in the case of birds.

It is well known that the productivity of an animal is not necessarily identical with its fecundity. However, to produce this much smaller quantity of eggs or of young, the sparrow and the rabbit I have chosen the most prolific bird and mammal expend a much larger quantity of material than the frog or the fish.

The sparrow and the rabbit employ in producing their progeny a bulk of material greater than the weight of their body, whilst the enormous quantity of eggs laid by the frog does not weigh more than one-seventh part of the body of the frog.

It may be laid down, as a general rule, that although fecundity, that is to say the number of eggs or of young which are produced, diminishes as the organism becomes more complex, the productivity on the other hand increases, expressed in percentage of weight.

The productivity, which is not more than 18 per cent, in batrachia, reaches 50 per cent, in reptiles, 74 per cent, in mammals, and 82 per cent, in birds.

It is plain that if reproduction shortens the life of mammals by weakening the organism, it must be the productivity, not the fecundity, which is the important factor.

I have just shown that productivity is greater in birds than in mammals, and in consequence it cannot be on account of any greater burden of reproduction that mammals have a shorter life than birds.

The shortness of mammalian life, again, cannot be attributed to the fact that mammals give birth to young, whilst the long-lived reptiles and birds produce eggs, because the longevity of the males, which produce neither young nor eggs, is none the less practically equal to that of the females of the same species.

The reason of the short life of mammals must be sought for elsewhere. No indication as to the cause of this difference can be found in the structure of the organs of circulation, respiration, or urinary secretion, or in the nervous or sexual apparatus.

The key to the problem is to be found in the organs of digestion. In reviewing the anatomical structure of the digestive apparatus in the vertebrate series, one soon comes to the striking fact that mammals are the only group in which the large intestine is much developed.

In fish, the large intes- tine is the least important part of the digestive tube, being little wider in calibre than the small intestine.

Amongst batrachia, where it is a relatively wide sack, it has begun to assume some importance. In several reptiles it is still larger, and may be provided with a lateral out-growth, which is to be regarded as a caecum.

In most birds, at the point where the large intestine passes into the small intestine, there is a pair of caeca, more or less developed.

These caeca are absent in climbing birds, such as the wood-pecker, the oriole, and many others. They are reduced to a pair of tiny out-growths in the eagles, sparrow-hawks, and other diurnal birds of prey, and in pigeons, and perching birds.

These organs are larger in the nocturnal birds of prey, in gallinaceous birds, and in ducks, etc.

Thus, for instance, in a rhea Rhea americana which I dissected, the caeca were nearly two-thirds as long as the small intestine.

The weight of the two caeca with their contents was more than 10 per cent, of the total weight of the bird. Notwithstanding the exceptions, which are relatively rare, the large intestine is badly developed in the case of birds.

On the other hand, it reaches its largest size amongst mammals. In these animals, " only the posterior portion of the latter, or rectum, which passes into the pelvic cavity, corresponds to the large intestine of lower Vertebrates; the remaining, and far larger part, must be looked upon as a neomorph, and is called the colon.

Maumus, " Les caecums des oiseaux," Annales des sciences naturel- les, See also P. Linnaan Soc. Newton Parker, p. Jeffrey Bell, B.

Owing ro its greater length, it is arranged in coils, so that the terminal portion only has the straight course taken by the hind-gut of other Vertebrata.

On the one hand mammals are shorter lived than birds and lower vertebrates, on the other hand the large intestine is much longer in them than in any other vertebrates.

Is there here any link of causality, binding the two Characters, or is it a mere coincidence? To answer the question we must turn to the function of the large intestine in vertebrates.

In the lower members of the group fish, batrachia, reptiles, birds, etc. It takes no share in digestion, as that is the function of the stomach and the small intes- tine.

Only the caecum can be thought to have some digestive property. In reptiles, the lowest vertebrates in which the caecum is present, it is so little differentiated from the large intestine itself, that it is difficult to assign to it any specialised function.

In very many birds, how- ever, the caeca are well separated from the main diges- tive tube. The food material passes into them in con- siderable quantities, and is retained there sufficiently long for some digestive process to take place.

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