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Filter by post date: 30 Apply filter Apply filter. Filter by nic geography:. Footer Navigation. According to him, it is theoretically obvious that a time must come when the replacement of the epidermic cells completely ceases.

As the superficial layers of the skin continue to dry up and be cast off, it is plain that the epidermis must disappear completely.

These theoretical considerations, however, are not com- patible with certain well-known facts indicating that there is no general cessation of the power of cell reproduction in old age.

The hairs and the nails, which are epidermic outgrowths, continue to grow throughout life, their growth being due to the proliferation of their constituent cells.

There is no sign of any arrest in the development of these structures, even in the most advanced old age. The reverse is true.

It is well known that the hairs on some parts of the body increase in number and in length in old people. In some lower races, for instance in the Mongols, the moustache and the beard grow vigorously in old age, whilst young people of the same race have only very small moustaches and practically no trace of beard.

So also in white women the fine and almost invisible down which covers the upper lip, the chin, and the cheeks in the young may become replaced by long hairs which form a mous- tache or beard.

Pohl, a specialist in the growth of hair, has measured the rate of growth in different circumstances. He has shown that in an old man of 61 the hair on the temple grew ii mm.

Plainly, there is no case here of a progressive diminution of cell-proliferation with age. 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.

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Dongguan Kecheng Communication Co. Hongan Group Co. Shenzhen Dowdon Technology Co. Shenzhen Focalvalue Technology Co. The caeca in birds.

The large intestine of mam- mals. Function of the large intestine. The intestinal microbes and their agency in producing auto-intoxication and auto-infection in the organism.

Passage of microbes through the intestinal wall Rumi- nants. The horse. Intestinal flora of birds. Intestinal flora of cursorial birds. Duration of life in cursorial birds.

Flying mammals. Intestinal flora and longevity of bats. Some exceptions to the rule. Resistance of the lower vertebrates to certain intestinal microbes Theory of Ebstein on the normal duration of human life.

Instances of human longevity. Examples of very old trees. Examples of short-lived plants. Pro- longation of the life of some plants.

Theory of the natural death of plants by exhaustion. Death of plants from auto- intoxication. Examples of natural death associated with violent acts.

Examples of natural death in animals without digestive organs. Natural death in the two sexes. Hypothesis as to the cause of natural death in animals.

Analogy of natural death and sleep. Theories of sleep. The instinct of sleep. The instinct of natural death.

Replies to critics. Agreeable sensation at the approach of death. Theory of " medical selection " as a cause of degeneration of the race.

Utility of prolonging human life The " immortality draught " of the Taoists. Brown-S6quard's method.

The spermine of Poehl. Weber's precepts. Increased duration of life in historical times. Hygienic maxims. Decrease in cutaneous cancer Prevention of syphilis.

Case of a woman whose large intestine was inactive for six months. Another case where the greater part of the large intestine was com- pletely shut off.

Attempts to disinfect the contents of the large intestine. Harmlessness of sterilised food. Means of preventing the putrefaction of food.

Lactic fermentation and its anti-putrescent action. Experiments on man and mice. Longevity in races which used soured milk. Comparative study of different soured milks.

Properties of the Bulgarian Bacillus. Actual existence of rudimentary organs. Atrophy of Jacobson's organ and of the Harderian gland in the human race.

Their muscular strength. Their expression of fear. The awakening of latent instincts of man under the influence of fear.

Natural somnambulism. Doubling of personality. Some examples of somnam- bulists. Analogy between somnambulism and the life of anthropoid apes.

The psychology of crowds. Importance of the investigation of hysteria for the problem of the origin of man Loss of individuality in the associations of lower animals.

Myxomycetes and Siphonophora. Individuality in Ascidians. Progress in the development of the individual living in a society.

Development and preservation of indi- viduality in colonies of insects. Division of labour and sacrifice of individuality in some insects Differentiation in the human race.

Learned women. Habits of a bee, Halictus quadricinctus. Col- lectivist theories. Progress of individuality in the societies of higher beings.

Pessimistic poets. Views of E. Analysis of Kowalev- sky's work on the psychology of pessimism History of a man of science who was pessimistic when young and who became an optimist in old age.

Optimism of Schopenhauer when old. Development of the sense of life. Development of the senses in blind people.

Pessimism of youth. Tendency to suicide. Work and love. His mode of life in that period. Influence of love in artistic production. Senile love of Goethe.

Relation between genius and the sexual activities Physical and intellectual vigour of the old man. Optimistic conception of life. The three monologues in the first Part.

Faust's pessimism. The brain-fatigue which finds a remedy in love. The romance with Mar- guerite and its unhappy ending.

Amorous passion of the old man. Humble attitude of the old Faust. Platonic love for Helena. The old Faust's conception of life. His optimism.

Vivisection and anti-vivi- section. Enquiry into the possibility of rational morality. Utilitarian and intuitive theories of morality.

Kant's theory of moral obligation. Some criticisms of the Kantian theory. History of two brothers brought up in the same circumstances, but whose conduct was quite different.

Late development of the sense of life. Evolution of sym- pathy. The sphere of egoism in moral conduct. Christian morality.

Morality of Herbert Spencer. Danger of exalted altruism. Com- parison with the modification of the constitution of plants and of animals. Schlanstedt rye.

Burbank's plants. The ideal of orthobiosis. The immorality of ignorance. The place of hygiene in the social life. The place of altru- ism in moral conduct.

Robineau, a lady of years of age Principal characters of old age Examples of old mammals Old birds and tor- toises Hypothesis of senile degeneration in the lower animals IN the ' ' Nature of Man ' ' I laid down the outlines of a theory of the actual changes which take place during the sen- escence of our body.

These ideas, on the one hand, have raised certain difficulties, and, on the other, have led to new investigations. As the study of old age is of great theoretical importance, and naturally is of practical value, I think that it is useful to pursue the subject still further.

In the Melanesian Islands, old people who have become incapable of doing useful work are buried alive. In times of famine, the natives of Tierra del Fuego kill and eat the old women before they touch their dogs.

Civilised races do not act like the Fuegians or other savages ; they neither kill nor eat the aged, but none the less life in old age often becomes very sad.

As they are incapable of performing any useful function in the family or in the village, the old people are regarded as a heavy burden.

Although they cannot be got rid of, their death is awaited with eagerness, and is never thought to come soon enough.

The Italians say that old women have seven lives. According to a Bergamask tradition, old women have seven souls, and after that an eighth soul, quite a little one, and after that again half a soul ; whilst the Lithuanians complain that the life of an old woman is so tough that it cannot be crushed even in a mill.

We may take it as an echo of such popular ideas that murders of old people are extremely common even in the most civilised European countries.

I have been astonished in looking through criminal records to see how many cases there are of the murder of old people, specially of old women.

It is easy to divine the motives of these acts. A convict of the Island of Saghalien, condemned for the assassination of several old persons, declared naively to the prison doctor : "Why pity them?

They were already old, and would have died in any case in a few years. In the middle of the conversation a student declares that he would " murder and rob any cursed old woman without the least remorse.

On the one hand a stupid old woman, childish, worthless, ill-tempered, and in bad health ; no one would miss her, indeed she is a nuisance to evervone.

On the other hand, there are fresh and vigorous young people who are dying in their thousands, in the most senseless way, no one troubling about them, and everywhere the same thing is going on.

They prefer death to a life oppressed by material hard- ships or burdened by diseases. The daily papers give many instances of old people who, tired of suffering, asphyxiate themselves by their charcoal stoves.

The frequency of suicide in the case of the old has been established by numerous statistics, and the new facts which I now cite do no more than confirm it.

In , in Prussia, amongst , individuals there were cases of suicide of men between the ages of 20 and 50, but , that is to say, nearly twice as many of men between the ages of 50 and In Denmark, a country in which suicide is notoriously common, a similar proportion exists.

Thus, in Copenhagen, in the ten years from to , there were suicides of men between 50 and These figures relate to , individuals.

In some countries laws have been passed to bring about this. For instance, a Danish law of June 27th, , established compulsory aid for the aged, enacting that every person more than 60 years old was to have the legal right to aid if required.

In more than 36, people 36, were pensioned 1 Westergaard, Mortalitaet u. Morbilitaet, 2nd. In Belgium, the indigent old people are not pensioned until they reach the age of In France, until recently, the aged poor could be supported at the public expense only by prosecut- ing them and sending them to prison for begging.

This state of affairs, however, ceased with the application of the law of July i5th, , according to which any French subject without resources, unable to support himself by work, and either more than 70 years of age, or suffering from some incurable infirmity or disease, is to receive public assistance.

It has been thought the proper course to make such laws, and to lay the burden on the general population, without inquiring if it may not be possible to retard the debility of old age to such an extent that very old people might still be able to earn their livelihood by work.

Old age can be studied by the methods of exact science, and there may yet be established some regimen by which health and vigour will be preserved beyond the age where now it is generally necessary to resort to public charity.

With this object, a systematic investigation of senescence should be made in institutions for the aged, where there are always a large number of people from 75 to 90 years old, although centenarians are extremely rare.

I know many institu- tions for aged men where, from their first foundation, there has been no case of an inhabitant reaching the age of , and even in similar institutions for women, although women live to much greater ages than men, centenarians are very rare.

At the Salpetriere, for instance, where there is always a large number of old women, it is the rarest chance to find a centenarian.

Opportunity for the study of the extremely aged is to be found only in private families. A few years ago an old woman who had reached her looth year was the pride of the Salpetriere.

She was bedridden and extremely feeble physically and mentally. She replied briefly when she was asked questions, but apparently with- out any idea of what they meant.

Not long ago, a lady who lived in a suburb of Rouen reached her looth birthday. The local newspapers wrote exaggerated articles about her, praising the integrity of her mind and her physical strength.

I paid a visit to her myself, hoping to make a detailed investigation, but I found at once that the journalists had completely misrepre- sented her condition.

Although her physical health was fairly good, her intelligence had degenerated to such an extent that I had to abandon the idea of any serious inves- tigation.

The most interesting of all the centenarians with whom I have become acquainted had reached an extremely advanced age, having entered upon her loyth year.

It is about two years ago that a journalist, Monsieur Flamans, took me to see this Mme. Robineau who lived in a suburb of Paris. I found her a very old-looking lady, rather short, thin, with a bent back, and leaning heavily on a cane when she walked.

The physical condition Mme. Robineau was born on January i2th, , of this woman of more than years, showed extreme decay.

She had only one tooth ; she had to sit down after every few steps, but, once comfortably seated, she could remain in that position for quite a long time.

She went to bed early and got up very late. Her features displayed very great age see Fig. Robineau, a centenarian. From a photograph taken on her one hundred and fifth birthday.

The skin of her hands had become so transparent that one could see the bones, the blood-vessels, and the tendons.

None the less, Dr. Lowenberg, a well-known aurist, had assured himself that her auditory organs showed in a most marked degree, the usual signs of old age, such as complete insensibility to high notes and slight deafness for low notes.

Lowenberg attributed these changes to senile degeneration of the ear which affected more and more seriously the nervous mechanism although it had caused little change in the conducting apparatus.

Notwithstand- ing her physical weakness, Mme. Robineau retained her intelligence fully, her mind remained delicate and refined and the goodness of her heart was touching.

In contrast with the usual selfishness of old people, Mme. Robineau took a vivid interest in those around her. Her conversation was intelligent, connected, and logical.

Examination of the physical functions of this old lady revealed facts of great interest. Ambard found that the sounds of the heart were normal, but perhaps a little accentuated.

The pulse was regular, 70 to 84 a minute, and its tension was normal. The arterial pressure was The lungs were sound. All these facts testify to her general health.

The most remarkable circumstance was the absence of sclerosis of the arteries, although such degeneration is usually believed to be a normal character of old age.

Analysis of the urine, made on several occasions, showed that the kidneys were affected with a chronic disease, which, however, was not serious.

There was no albumen or sugar. The sediment contained crystals of uric acid, some pavement epithelium cells, a very few cells from the tubules, some hyaline platelets and isolated white corpuscles.

She ate and drank little, "but her diet was varied. She took butcher's meat or chicken ex- tremely seldom, but ate eggs, fish, farinaceous food, vege- tables, and stewed fruit, and drank sweetened water with a little white wine, and sometimes, after a meal, a small glass of dessert wine.

The processes of alimentary diges- tion and excretion were normal. It has sometimes been thought that duration of life is a hereditary property.

There was no evidence for this in the present case. Madame Robineau's relatives had died comparatively early in life, and a centenarian was unknown in her family.

Her great age was an acquired character. Her whole life had been extremely regular. She had married a timber merchant, and had lived for many years in a suburb of Paris in comfortable circum- stances.

Her character was gentle and affectionate; she was thoroughly domesticated, and had been devoted to home life with very few distractions.

At the age of years, her intelligence suddenly became weak. She lost her memory almost completely, and some- times wandered.

But her gentle and affectionate disposi- tion remained unaltered. The appearance of aged persons is too well known to make detailed description necessary.

The skin of the face is dry and wrinkled and generally pale; the hairs on the head and the body are white; the back is bent, and the gait is slow and laborious, whilst the memory is weak.

Such are the most familiar traits of old age. Baldness is not a special character; it often begins during youth and naturally is progressive, but if it has not already appeared, it does not come on with old age.

The stature diminishes in old age. In extreme cases, the loss may be nearly three inches. The weight also becomes less.

According to Quetelet, males attain their maximum weights at the age of forty, females at that of fifty. From the age of sixty years onwards, the body becomes lighter, the loss at eighty being as much as thirteen pounds.

Such losses of height and Weight are signs of the general atrophy of the aged organism. Not merely the soft parts, such as the muscles and viscera, but even the bones lose weight, in the latter case the loss being of the mineral constituents.

This process of decalcification makes the skeleton Brittle, and is sometimes the cause of fatal acci- dents.

The v61ume diminishes, and the substance becomes paler; the fat between the fibres is absorbed, and may disappear completely. Movements are slower, and the muscular force is abated.

This progressive degeneration has been examined by dynamometrical measurements of the hand and the trunk, and is greater in males than in females.

The volumes and weights of the visceral organs simi- larly become smaller, but the diminution is not uniform.

The old age of lower mammals presents characters similar to those found in man. I can now give other instances than the case of the old dog which I described in the " Nature of Man.

The eyes are usually dim, and discharge an abnormal quantity of water. The margin of the ears, specially on the lower side, is usually frayed.

The skin of the trunk is rough- ened, hard, and warty, so that the organ has lost much of its flexibility.

The skin on the body generally is worn and wrinkled; the legs are thinner than in maturity, the FIG.

A Mare, thirty-seven years old. The skin round the toe-nails is roughened and frayed. The tail is scaly and hard, and the tip is often hairless.

Horses begin to grow old much sooner than elephants. I reproduce Fig. The skin, bare in places, but elsewhere covered with long hairs, shows considerable atrophy.

The general attitude reveals the feebleness of the whole body. Many birds, on the other hand, show at similar ages very slight external change, as may be seen from the photograph of a duck more than 25 years old Fig.

Jean Charcot. At a still greater age, as may be seen occasionally in parrots, FIG. A White Duck, which lived for more than a quarter of a century.

On the other hand, the oldest reptiles which have been observed do not differ in appearance from normal adults of the same species.

I have in my posses- sion a male tortoise Testudo mauritanica given me by my friends MM. Rabaud and Caullery, and which is at least 86 years old.

More than 31 years ago it was wounded by a blow, the traces of which remain visible on the right side of the carapace Fig. In the last three years the tortoise lived in a garden at Montauban, along with two females which laid fertile eggs.

The old male, although, as I have said, probably at least 86 years of age, was still sexually healthy. I have borrowed from the interesting volume of Prof.

Sir E. Ray Lankester 1 the figure Fig. An Old Land-tortoise. It was brought to Mauritius from the Seychelles in , and has lived since then in the garden of the Governor, and as it has thus already been years in captivity, its age must be at least years, although we have not exact information.

Notwithstanding this, it shows no signs of old age. The examples which I have brought together show that 1 Extinct Animals, London, , pp.

I think it a fair inference that senility, the precocious senescence which is one of the greatest sorrows of humanity, is not so profoundly seated in the constitution of the higher animals as has generally been supposed.

It is not necessary, therefore, to discuss at FIG. A Water-tortoise, mere than years old. After Prof. Ray Lankester.

I have already shown, in the " Nature of Man," the differ- ence which exists between senile degeneration in our own bodies and the phenomena of senescence amongst In- fusoria which, as M.

Maupas described, are followed by a process of rejuvenescence. According to the more recent results of several investigators, the difference is still greater than I had supposed.

Enriquez l has been able 1 Rendiconti d. Lincei, , vol xiv. Here we are far from the condition in the human race. Hertwig, 1 one of the best observers of the lower animals, has recently attempted to show that the very simple animalculae of the genus Actinosphaerium are subject to true physiological degeneration.

He has several times seen cultures of this Rhizopod degenerate, until all the individuals had died, notwithstanding the presence of abundant food.

Hertwig attributed this to the " con- stitution of the Actinosphaerium having been weakened by too great vital activity at an earlier stage.

As this idea had not occurred to the observer, he had not searched for parasitic microbes amongst the granulations which are always present in the body of an Actinosphaerium.

How- ever this may be, I cannot accept the facts brought forward by this distinguished German as a valid proof of the exist- ence of senile degeneration in these lowly creatures.

The facts that I have brought together in this chapter justify the conclusion that human beings who reach ex- treme old age may preserve their mental qualities notwith- standing serious physical decay.

Moreover, it is equally plain that the organism of some vertebrates is- able to resist the influence of time much longer than is the case with man under present conditions.

Jena, Marinesco, the neurono- phags are true phagocytes The whitening of hair and the destruction of nerve cells, as arguments against a theory of old age based on the failure of the reproductive powers of the cells ALTHOUGH it has not been proved that living matter must inevitably undergo senile decrepitude, it is none the less true that man and his nearest allies generally exhibit such degeneration.

It is therefore extremely important to recog- nise the real causes of our senescence. There have been many hypotheses on the subject, but there are compara- tively few definite facts known.

Biitschli has supposed that the life of cells is maintained by a specific vital ferment which becomes feebler in pro- portion to the extent of cellular reproduction, but I cannot regard this as more than a pious opinion.

The ferment has never been seen, and we do not know of its actual existence. According to the better-known theory of Prof.

As old age appears at different times in different species and different indivi- duals, Weismann has concluded that the possible number of cell generations differs in different cases.

He has not found, however, a solution of the problem as to why multi- plication of cells should cease in one individual, whereas it proceeds much further in other individuals.

Minot, 1 the American zoologist, has developed a similar theory, and has employed an exact method to determine the gradual diminution in the rate of growth of an animal from its birth onwards.

According to him, the power of reproduction of the cells weakens progressively during life, until a point is necessarily reached at which the organism, no longer capable of repairing itself, begins to atrophy and degenerate.

Buehler 2 has recently laid stress upon this theory. There is no doubt that cells reproduce much more actively during the embryonic period.

The process becomes slower later on, but, none the less, continues to display itself throughout the whole period of life. Buehler attributes the difficulty with which certain wounds heal in the case of old people to the insufficiency of cellular reproduc- tion.

He thinks in particular that the proliferation of the cells of the skin, to replace those which are worn off from the surface, becomes less active with age.

According to him, it is theoretically obvious that a time must come when the replacement of the epidermic cells completely ceases.

As the superficial layers of the skin continue to dry up and be cast off, it is plain that the epidermis must disappear completely.

These theoretical considerations, however, are not com- patible with certain well-known facts indicating that there is no general cessation of the power of cell reproduction in old age.

The hairs and the nails, which are epidermic outgrowths, continue to grow throughout life, their growth being due to the proliferation of their constituent cells.

There is no sign of any arrest in the development of these structures, even in the most advanced old age.

The reverse is true. It is well known that the hairs on some parts of the body increase in number and in length in old people. In some lower races, for instance in the Mongols, the moustache and the beard grow vigorously in old age, whilst young people of the same race have only very small moustaches and practically no trace of beard.

So also in white women the fine and almost invisible down which covers the upper lip, the chin, and the cheeks in the young may become replaced by long hairs which form a mous- tache or beard.

Pohl, a specialist in the growth of hair, has measured the rate of growth in different circumstances.

He has shown that in an old man of 61 the hair on the temple grew ii mm. Plainly, there is no case here of a progressive diminution of cell-proliferation with age.

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.

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