Transcriber’s Note:
The position of the footnote anchor 171 at page 229 is a guess of the transcriber as the anchor was missing in the original book.
The book cover image was created by the transcriber and is placed in the public domain.

THE PROLONGATION OF LIFE

OPTIMISTIC STUDIES

BY

ÉLIE METCHNIKOFF

SUB-DIRECTOR OF THE PASTEUR INSTITUTE, PARIS

THE ENGLISH TRANSLATION

EDITED BY

P. CHALMERS MITCHELL
M.A., D.SC. OXON., HON. LL.D., F.R.S.
Secretary of the Zoological Society of London; Corresponding Member
of the Academy of Natural Sciences of Philadelphia

G. P. PUTNAM’S SONS
NEW YORK & LONDON
The Knickerbocker Press
1908

EDITOR’S INTRODUCTION

Élie Metchnikoff has carried on the high purpose of the Pasteur Institute by devoting his genius for biological inquiry to the service of man. Some years ago, in a series of Essays which were intended to be provocative and educational, rather than expository, he described the direction towards which he was pressing. I had the privilege of introducing these Essays to English readers under the title The Nature of Man, a Study in Optimistic Philosophy. In that volume, Professor Metchnikoff recounted how sentient man, regarding his lot in the world, had found it evil. Philosophy and literature, religion and folk-lore, in ancient and modern times have been deeply tinged with pessimism. The source of these gloomy views lies in the nature of man itself. Man has inherited a constitution from remote animal ancestors, and every part of his structure, physical, mental and emotional, is a complex legacy of diverse elements. Possibly at one time each quality had its purpose as an adaptation to environment, but, as man, in the course of his evolution, and the environment itself have changed, the old harmonious intercourse between quality and circumstances has been dislocated in many cases. And so there have come into existence many instances of what the Professor calls “disharmony,” persistences of structures, or habits, or desires that are no longer useful, but even harmful, failures of parallelism between the growth, maturity and decay of physical and mental qualities and so forth. Religions and philosophies alike have failed to find remedies or efficient anodynes for these evils of existence, and, so far, man is justified of his historical and actual pessimism.

Metchnikoff, however, was able to proclaim himself an optimist, and found, in biological science, for the present generation a hope, or, at the least, an end towards which to work, and for future generations a possible achievement of that hope. Three chief evils that hang over us are disease, old age, and death. Modern science has already made vast strides towards the destruction of disease, and no one has more right to be listened to than a leader of the Pasteur Institute when he asserts his confidence that rational hygiene and preventive measures will ultimately rid mankind of disease. The scientific investigation of old age shows that senility is nearly always precocious and that its disabilities and miseries are for the most part due to preventable causes. Metchnikoff showed years ago that there exists in the human body a number of cells known generally as phagocytes, the chief function of which is to devour intruding microbes. But these guardians of the body may turn into its deadly enemies by destroying and replacing the higher elements, the specific cells of the different tissues. The physical mechanism of senility appears to be in large measure the result of this process. Certain substances, notably the poisons of such diseases as syphilis and the products of intestinal putrefaction, stimulate the activity of the phagocytes and so encourage their encroachment on the higher tissues. The first business of science is to remove these handicaps in favour of the wandering, corroding phagocytes. Specific poisons must be dealt with separately, by prevention or treatment, and it is well known that Metchnikoff has made great advances in that direction. The most striking practical side of The Nature of Man, however, was the discussion of the cause and prevention of intestinal putrefaction. Metchnikoff believes that the inherited structure of the human large intestine and the customary diet of civilised man are specially favourable to the multiplication of a large number of microbes that cause putrefaction. The avoidance of alcohol and the rigid exclusion from diet of foods that favour putrefaction, such as rich meats, and of raw or badly cooked substances containing microbes, do much to remedy the evils. But the special introduction of the microbes which cause lactic fermentation has the effect of inhibiting putrefaction. By such measures Metchnikoff believes that life will be greatly prolonged and that the chief evils of senility will be avoided. It may take many generations before the final result is attained, but, in the meantime, great amelioration is possible. There remains the last enemy, death. Metchnikoff shows that in the vast majority of cases death is not “natural,” but comes from accidental and preventable causes. When diseases have been suppressed and the course of life regulated by scientific hygiene, it is probable that death would come only at an extreme old age. Metchnikoff thinks that there is evidence enough at least to suggest that when death comes in its natural place at the end of the normal cycle of life, it would be robbed of its terrors and be accepted as gratefully as any other part of the cycle of life. He thinks, in fact, that the instinct of life would be replaced by an instinct of death.

Metchnikoff’s suggestion, then, was that science should be encouraged and helped in every possible way in its task of removing the diseases and habits that now prevent human life from running its normal course, and his belief is that were the task accomplished, the great causes of pessimism would disappear.

In this new volume, The Prolongation of Life, the main thesis is carried further, and a number of criticisms and objections are met. The latter, so far as they relate to technical details, I need say nothing of here, as Metchnikoff and his staff at the Pasteur Institute are the most skilled existing technical experts on these matters, but I cannot refrain from a word of comment on the brilliant treatment of the objection to the suggested amelioration of human life that it considered only the individual and neglected the just subordination of the individual to society. In the sixth Part of this volume, Metchnikoff discusses the relation of the individual to the species, society or colony, from the general point of view of comparative biology, and shows that as organisation progresses, the integrity of the individual becomes increasingly important. Were orthobiosis, the normal cycle of life, attained by human beings, there still would be room for specialisation of individuals and for differentiation of the functions of individuals in society, but instead of the specialisation and differentiation making individuals incomplete throughout their whole lives, they would be distributed over the different periods of the life of each individual.

As these lines are intended to be an introduction, not a commentary, I will now leave the reader to follow the argument in the book itself.

P. Chalmers Mitchell.

London, August, 1907.

PREFACE

It is now four years since I wrote a volume, the English translation of which was called The Nature of Man, and which was an attempt to frame an optimistic conception of life. Human nature contains many very complex elements, due to its animal ancestry, and amongst these there are some disharmonies to which our misfortunes are due, but also elements which afford the promise of a happier human life.

My views have encountered many objections, and I wish to reply to some of these by developing my arguments. This was my first task in this book, but I have also brought together a series of studies on problems which closely affect my theory.

Although it has been possible to support my conception by new facts, some of which have been established by my fellow-workers, others by myself, there still remain many sides of the subject where it is necessary to fall back on hypotheses. I have accepted such imperfections instead of delaying the publication of my book.

Even at present there are critics who regard me as incapable of sane and logical reasoning. The longer I postpone publication, the longer would I leave the field open to such persons. What I have been saying may serve also as a reply to the remark of one of my critics, that my ideas have been “suggested by self-preoccupation.”

It is, of course, quite natural that a biologist whose attention had been aroused by noticing in his own case the phenomena of precocious old age should turn to study the causes of it. But it is equally plain that such a study could give no hope of resisting the decay of an organism which had already for many years been growing old. If the ideas which have come out of my work bring about some modification in the onset of old age, the advantage can be gained only by those who are still young, and who will be at the pains to follow the new knowledge. This volume, in fact, like my earlier one on the “Nature of Man,” is directed much more to the new generation than to that which has already been subjected to the influence of the factors which produce precocious old age. I think that thus the experience of those who have lived and worked for long can be made of service to others.

As this volume is a sequel to The Nature of Man, I have tried as much as possible to avoid repetition of what was fully explained in the earlier volume.

Here I bring together the results of work that has been done since the publication of The Nature of Man. Some of the chapters relate to subjects upon which I have lectured, or which, in a different form, have been printed before. For instance, the section on the psychic rudiments of man appeared in the Bulletin de l’Institut général psychologique of 1904, the essay on Animal Societies was published in the Revue Philomatique de Bordeaux et du Sud-Ouest of 1904, and in the Revue of J. Finot of the same year, whilst a German translation of it appeared in Prof. Ostwald’s Annalen der Naturphilosophie. The chapter on soured milk first appeared as a pamphlet, published in 1905. The substance of my views on natural death was published in June last in “Harper’s Monthly Magazine” of New York, while the chapter on natural death in animals appeared in the first number of the Revue du Mois for 1906.

I have to thank most sincerely the friends and pupils who have helped me by bringing before me new facts, or other materials; the names of these will appear in their proper places in the volume. I have not mentioned by name, however, Dr. J. Goldschmidt, whose continual encouragement and practical sympathy have made my work much easier.

Finally, my special thanks are due to Drs. Em. Roux and Burnet, and M. Mesnil, who have been so good as to correct my manuscript and the proofs of this volume.

É. M.

Paris, Feb. 7, 1907.

CONTENTS

THE PROLONGATION OF LIFE

PART I
THE INVESTIGATION OF OLD AGE

I
THE PROBLEMS OF SENILITY

Treatment of old people in uncivilised countries—Assassination of old people in civilised countries—Suicide of old people—Public assistance in old age—Centenarians—Mme. Robineau, a lady of 106 years of age—Principal characters of old age—Examples of old mammals—Old birds and tortoises—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 senescence 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.

Although there exist races which solve the difficulty of old age by the simple means of destroying aged people, the problem in civilised countries is complicated by our more refined feelings and by considerations of a general nature.

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. When they were asked why they did this, they said that dogs could catch seals, whilst old women could not do so.

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 naïvely to the prison doctor: “Why pity them? They were already old, and would have died in any case in a few years.”

In the celebrated novel of Dostoiewsky, “Crime and Punishment,” there is a tavern scene where young people discuss all sorts of general topics. In the middle of the conversation a student declares that he would “murder and rob any cursed old woman without the least remorse.” “If the truth were told,” he goes on to say, “this is how I look at the thing. 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 everyone. She does not even herself know any reason why she should live, and perhaps to-morrow death will make a good riddance of her. 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.”

Old people not only run the risk of murder; they very often end their own lives prematurely by suicide.

They prefer death to a life oppressed by material hardships 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 1878, in Prussia, amongst 100,000 individuals there were 154 cases of suicide of men between the ages of 20 and 50, but 295, that is to say, nearly twice as many of men between the ages of 50 and 80. In Denmark, a country in which suicide is notoriously common, a similar proportion exists. Thus, in Copenhagen, in the ten years from 1886 to 1895, there were 394 suicides of men between 50 and 70. These figures relate to 100,000 individuals. Of the suicides 36-1/2 per cent. were those of people in the prime of life, 63-1/2 per cent. those of the aged.[1]

In such circumstances, it is natural that politicians and philanthropists have made many attempts to ameliorate the old age of the poor. In some countries laws have been passed to bring about this. For instance, a Danish law of June 27th, 1891, 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 1896 more than 36,000 people (36,246) were pensioned under this law, at a cost of nearly £200,000. In Belgium, the indigent old people are not pensioned until they reach the age of 65. In France, until recently, the aged poor could be supported at the public expense only by prosecuting them and sending them to prison for begging. This state of affairs, however, ceased with the application of the law of July 15th, 1905, 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 institutions for aged men where, from their first foundation, there has been no case of an inhabitant reaching the age of 100, and even in similar institutions for women, although women live to much greater ages than men, centenarians are very rare. At the Salpêtrière, 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.

Most of the centenarians whom I have been able to see have been so defective mentally that all that can be studied in them are the physical qualities and functions. A few years ago an old woman who had reached her 100th year was the pride of the Salpêtrière. She was bedridden and extremely feeble physically and mentally. She replied briefly when she was asked questions, but apparently without any idea of what they meant.

Not long ago, a lady who lived in a suburb of Rouen reached her 100th 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 misrepresented 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 investigation.

The most interesting of all the centenarians with whom I have become acquainted had reached an extremely advanced age, having entered upon her 107th 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 12th, 1800), of this woman of more than 106 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. [1]), although her skin was not extremely wrinkled.

Fig. 1.—Mme. 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. Her senses were very feeble; she could see only with one eye; taste and smell were extremely rudimentary; her hearing was her best means of relation with the external world. None the less, Dr. Löwenberg, 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. Dr. Löwenberg 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. Notwithstanding 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. Dr. 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 17. 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.[2]

Although the sense of taste was weak, Madame Robineau had a fair appetite. She ate and drank little, but her diet was varied. She took butcher’s meat or chicken extremely seldom, but ate eggs, fish, farinaceous food, vegetables, 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 digestion 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 circumstances. 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 106 years, her intelligence suddenly became weak. She lost her memory almost completely, and sometimes wandered. But her gentle and affectionate disposition 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. As the result of a series of observations, it has been established that a man loses more than an inch (3·166 cm.), and a woman more than an inch and a half (4·3 cm.), between the ages of fifty and eighty-five years. In extreme cases, the loss may be nearly three inches. The weight also becomes less. According to Quételet, 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 accidents.

The loss of muscular tissue is specially great. The volume 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 similarly 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.”

I will first take the case of old elephants, described by a competent observer. “The general appearance is wretched, the skull being often hardly covered with skin; there are deep abrasions under the eyes, and smaller ones on the cheeks, whilst the skin of the forehead is very often deeply fissured or covered with lumps. 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 roughened, 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 huge mass of muscles being much shrunken, whilst the circumference, especially just above the feet, is considerably reduced. The skin round the toe-nails is roughened and frayed. The tail is scaly and hard, and the tip is often hairless.

Fig. 2.—A Mare, thirty-seven years old.

Fig. 3.—A White Duck, which lived for more than a quarter of a century.

Horses begin to grow old much sooner than elephants. I reproduce (Fig. [2]) the photograph of a rare instance of longevity, a mare 37 years old, which belonged to M. Métaine, in the department of Mayenne. 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. [3]) which belonged to Dr. Jean Charcot. At a still greater age, as may be seen occasionally in parrots, the general debility of the body reveals itself in the attitude, in the condition of the feathers, and in the swelling of the joints. 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 possession a male tortoise (Testudo mauritanica) given me by my friends MM. Rabaud and Caullery, and which is at least 86 years old. It shows no sign of old age, and in all respects behaves like any other individual of this species. 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. [4]). 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.

Fig. 4.—An Old Land-tortoise.

I have borrowed from the interesting volume of Prof. Sir E. Ray Lankester[3] the figure (Fig. [5]) and description of a giant tortoise from the island of Mauritius, which is probably the oldest of all living animals. It was brought to Mauritius from the Seychelles in 1764, and has lived since then in the garden of the Governor, and as it has thus already been 140 years in captivity, its age must be at least 150 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 often amongst vertebrates there are some animals the organisms of which withstand the ravages of time much better than that of man. 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 length the general question as to whether senile degeneration is an inevitable event in living organisms.

Fig. 5.—A Water-tortoise, more than 150 years old.
(After Prof. Sir E. Ray Lankester.)

I have already shown, in the “Nature of Man,” the difference which exists between senile degeneration in our own bodies and the phenomena of senescence amongst Infusoria 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[4] has been able to propagate Infusoria to the 700th generation without any sign of senility being displayed. Here we are far from the condition in the human race.

R. Hertwig,[5] one of the best observers of the lower animals, has recently attempted to show that the very simple animalculæ of the genus Actinosphærium 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. Prof. Hertwig attributed this to the “constitution of the Actinosphærium having been weakened by too great vital activity at an earlier stage.” I should have thought that it was a much more natural explanation to suppose that the culture had undergone infection by one of the contagious diseases which so often destroy cultures of different kinds of lower animals and plants. 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 Actinosphærium. However this may be, I cannot accept the facts brought forward by this distinguished German as a valid proof of the existence 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 extreme old age may preserve their mental qualities notwithstanding 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.

II
THEORIES OF CAUSATION OF SENILITY

Hypothesis of the causation of senility—Senility cannot be attributed to the cessation of the power of reproduction of the cells of the body—Growth of the hair and the nails in old age—Inner mechanism of the senescence of the tissues—Notwithstanding the criticisms of M. Marinesco, the neuronophags 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 recognise the real causes of our senescence. There have been many hypotheses on the subject, but there are comparatively few definite facts known.

Bütschli has supposed that the life of cells is maintained by a specific vital ferment which becomes feebler in proportion 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. Weismann, old age depends on a limitation in the power of cells to reproduce, so that a time comes when the body can no longer replace the wastage of cells which is an inevitable accompaniment of life. As old age appears at different times in different species and different individuals, 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 multiplication of cells should cease in one individual, whereas it proceeds much further in other individuals. Prof. Minot,[6] 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. Dr. Buehler[7] 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 reproduction. 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. Buehler thinks that there must be a similar fate for the genital glands, the muscles, and all the other organs.

These theoretical considerations, however, are not compatible 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 moustache or beard.

Dr. 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 11 mm. in a month; on the other hand, the hair on the same region in boys of 11 to 15 years old grew in the same time only from 11 to 12 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. a month, whilst in the same individuals, at the age of 61 years, the rate of growth was only 11 mm.; but this diminution in the rate of growth is only apparent. The first figure concerned the hair taken from different regions of the scalp, whilst the second related only to the hair on the temples, and Dr. 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., and often less even than the 11 mm. recorded in the old man of 61.

I have been able to note that the nails grow even in very old people. In the case of Mme. Robineau, the centenarian, the nail of the middle finger of the left hand grew 2-1/2 mm. in three weeks. In the case of a lady of 32 years old, the corresponding nail grew 3 mm. in two weeks, the difference being out of all proportion to the enormous difference in the age. 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 criticism, especially with regard to the part played by the macrophags in the senescence of nervous tissue.

Neurologists in particular, have criticised my interpretation. For several years M. Marinesco[8] 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 preparations made from the brains of two very old persons. After careful examination I was convinced that my opponent had been inexact. In the brain of the two centenarians (one of whom died at the age of 117 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 difficult 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.”

Without taking notice of my reply, M. Marinesco has published another criticism of my theory in an article[9] entitled “Histological Investigations into the Mechanism of Senility.” In that work, although he himself had invented the designation “neuronophag” for a phagocyte that devours nerve-cells, he denies the existence of such a power. He thinks that nerve-cells atrophy independently of the cells that surround them. The latter, the so-called neuronophags, 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. There is no question of phagocytosis, of the existence of cells that devour their neighbours.

M. Léri has taken a similar view in a Report on the Senile Brain[10] 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.” In his monograph “La Neuronophagie,”[11] M. 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. 86).” Still more recently MM. Laignel-Lavastine and Voisin[12] have taken the same view, maintaining that the neuronophags 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 alterations 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 neuronophags in their preparations had been subjected to violent treatment. When M. Léri 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 observing cells destroyed by the processes of the laboratory. The illustrations in the memoir of M. 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 protoplasm; 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 particular preparations, had a false idea of its appearance.

In well-made preparations, neuronophags are typical cells with abundant protoplasm. When they have been preserved by a process that does not dissolve their contents, they show granules like those found in nerve-cells.

To study neuronophagy, M. Manouélian,[13] in the laboratory of the Pasteur Institute in Paris, set himself to improve the technical methods of preparation. 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. “My observations on the cerebro-spinal ganglia of human cases of hydrophobia,” he wrote, “show clearly that the macrophags act as phagocytes of the nerve-cells.” “Most of the cells in the nerve-ganglia contain yellow, brown, and black pigmented granules, usually united in small masses. What becomes of these granulations on the destruction and disappearance 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.”

By the aid of a very delicate mode of preparation, M. Manouélian has shown that in the case of senile brains the granules of the nerve-cells are absorbed by neuronophags. I have myself studied M. Manouélian’s preparations and can testify to the accuracy of his observations (Figs. [6] and [7]).

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 gradually 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. As I cannot state definitely what are the conditions that induce the phagocytosis of nerve-cells, I shall not attempt a discussion of the problem.

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. 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.

The general result of my investigation into the criticisms that have been published on this matter has confirmed me in my belief that neuronophagy plays a most important part in senescence, and recent observations that I have made with M. 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 reproductive 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.

III
MECHANISM OF SENILITY

Action of the macrophags in destroying the higher cells—Senile degeneration of muscular fibres—Atrophy of the skeleton—Atheroma and arterial sclerosis—Theory that old age is due to alteration in the vascular glands—Organic tissues that resist phagocytosis

The instances which I have selected in attempting to describe the mechanism of senescence of the tissues are not the only cases in which the importance of phagocytosis is evident. The blanching of hair is due to the destructive agency of chromophags; in atrophy of the brain neuronophags 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 infectious diseases, partly because it is the method of macrophags 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. [8]), where the surrounding macrophags gradually seize hold of the granules within it and carry these off. As the process goes on, the ovum becomes reduced to a shapeless mass, and finally leaves only a few fragments, or disappears completely. M. Matchinsky[14] 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.

Fig. 8.—Ovum of a Bitch in process of destruction by Phagocytes, which are full of fatty granules.
(After M. Matchinsky.)

The phenomena of atrophy in general and of senile decay afford other cases of tissue destruction in which the phagocytic character of the process is more modified and obscure than in nerve-cells and ova.

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 unimpaired admit their muscular weakness. The physical correlate of this condition is an actual atrophy of the muscles, and has for long been known to observers. More than half a century ago, Kölliker,[15] 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 frequently arranged in longitudinal series and present such signs of active division as are found in embryonic tissue.”

Other investigators afterwards made similar observations. Vulpian[16] and Douaud[17] 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 cases of muscular atrophy in old human beings and lower animals. We were able to recognise the phenomena observed by our predecessors. In senile atrophy the muscular fibres contain many nuclei, and these, increasing rapidly, bring about an almost complete disappearance of the contractile substance (Fig. [9]). The fibres preserve their striation for a certain time but eventually lose it and appear to contain an amorphous mass with numerous, rapidly multiplying nuclei.

Fig. 9.—Degeneration of striated muscle Fibres from the auricular muscle of a man aged 87 years.
(From a preparation made by Dr. Weinberg.)

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 proliferation, although the latter has frequently been suggested as the mechanism of senescence. 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 multiply. Just as in the atrophy of the brain there is an increase in the volume of neuroglœa, 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 sarcoplasm. The latter replaces the myoplasm, the specific striated substance of muscles, by a process which must be regarded as parallel with phagocytosis. 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 phagocytically with regard to the myoplasm, just as the chromophag 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 skeleton, 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, however, by the agency of cells very like some of the macrophags. These cells contain many nuclei, and are known as osteoclasts. They form round about the bony lamellæ and lead to their destruction, but are incapable of breaking off fragments of bone and dissolving them in their interiors. Although the intimate mechanism of this destructive action is not thoroughly understood, it seems probable that the cells secrete some acid which softens bone by dissolving the lime salts. 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. [10].

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-vertebrate discs in particular becoming impregnated with salts, so that the well-known senile malformation of the backbone is produced.

Fig. 10.—Destruction by osteoclasts of bony matter in the sternum of a man aged 81 years.
(From a preparation made by Dr. Weinberg.)

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 degeneration, 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 between the two alterations are very evident in the skull; the meningeal artery becomes sinuous and atheromatous, and the grooves on the inner side of the bones of the skull in which it runs, flatten out, and become larger because of other malformations.[18]

There is no disharmony in the nature of old people so striking as this transference of the lime salts from the skeleton to the blood-vessels, producing as it does a dangerous softening of the former, and a hardening of the latter that interferes with their function of carrying nutrition to the organs. 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 investigations 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 inflammatory 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 methods have not succeeded, but M. Josué[19] has been able to produce true arterial atheroma in rabbits by injecting into them adrenaline, the secretion of the supra-renal capsules.

This experiment has been repeated many times and is now well known. Later on, M. Boveri[20] 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 inflammations 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 supra-renal 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. Dr. Lorand[21] 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.” It has long been noticed that persons affected with myxodema, as a result of the degeneration of the thyroid gland, look like very old people. 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 profound bodily changes, is the result of degeneration of the thyroid gland. On the other hand, it is well known that in old people the thyroid and the supra-renals frequently show cystic degeneration. 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, however, that their action in producing senility is exclusive, or even preponderating. M. Weinberg, at the Pasteur Institute, made special investigations on this point, and found that the thyroid gland and the supra-renal 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 pneumonia, 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 result that what is due to infection has been set down as a symptom of old age.[22]

Although the appearance of patients from whom the thyroid gland has been removed, or in whom it has degenerated spontaneously, recalls that of old people, it is possible to exaggerate the similarity. In the masterly accounts of such unfortunates, recently compiled by the well-known surgeon Kocher[23] there are many points which are characteristic, without being typical, of old people.

Oedema of the skin which characterises thyroid patients is by no means usual in old age. The loss of hair, normal in the patients, is not a character of old age. In myxedematous 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 existence of any strong affinity between old age and affection of the thyroid gland. It is known that removal of the thyroid is followed by cachexia only in young subjects, MM. Bourneville and Bricon[24] 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 rare.

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[25] 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 information 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 they are destroyers of poisons, I cannot agree with the theory of Dr. Lorand.

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.

Fig. 11.—Testis tissue from a dog aged twenty-two years.
(From a preparation made by Dr. Weinberg.) I have already given an example (“The Nature of Man,” p. 98) of an old man of 94 in whom active spermatozoa were produced. I know of a similar case, the age being 103 years. Such cases are not rare, and not only in old men, but in old animals, the testes continue to be active. Dr. Weinberg and I have investigated these organs in a dog which died at the age of 22 years after several years of pronounced senility. 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. [11]). 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. None the less, this dog although markedly senile (Fig. [12]) still showed sexual instincts until shortly before it died.

Fig. 12.—An old dog, aged eighteen years.

It is manifest that the tissues do not invariably degenerate 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 degeneration but remain sufficiently active to produce macrophags which destroy the higher cellular elements of the body. I have frequently noticed cell division in such organs, and as an example may give the case of the bone marrow taken from a man of 81 years (Fig. [13]).

The eye is an organ that is modified in old age without the action of macrophags. Cataract and the senile arc which appears as a milky ring at the edge of the cornea are frequent in old age. These modifications are due to impregnation of the parts affected by fatty matter which makes them opaque. This deposition of fat[26] 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 consequence for anatomical reasons. Most organs possess in addition to their higher elements a constant source of macrophags. Such a source of phagocytosis is the neuroglœa

Fig. 13.—Bone marrow from the sternum of a man aged eighty-one years.
(From a preparation made by Dr. Weinberg.) in nervous tissues, the sarcoplasm in muscular tissues; the bones contain osteoclasts 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.

Some infectious diseases bring about precocious senility. A syphilitic child is “a miniature old man, with wrinkled face, skin dull and discoloured and flabby and hanging in folds as if it were too large.”[27] In such a case the active agent is the microbe of syphilis which has poisoned the child on the breast of its mother. 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, amongst other changes, there is deposition of fatty matter. The phagocytes resist the influence of invading poisons better than any of the other cells of the body and sometimes 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.

PART II
LONGEVITY IN THE ANIMAL KINGDOM

I
THEORIES OF LONGEVITY

Relation between longevity and size—Longevity and the period of growth—Longevity and the doubling in weight after birth—Longevity and rate of reproduction—Probable relation between longevity and the nature of the food

The duration of the life of animals varies within very wide limits. Some, as for instance, the males of certain wheel animalculæ (Rotifera) complete their cycle of life from birth to death in 50 or 60 hours, whilst others, like some reptiles, live more than 100 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.

As a general rule it may be said that a large animal takes more time than a small one to reach maturity, and it has been inferred from this that the length of the periods of gestation and of growth were in proportion to the longevity. Buffon[28] long ago stated his opinion that the “total duration of life bore some definite relation to the length of the period of growth.” Therefore, as the period of growth is, so to say, inherent in the species, longevity would have to be regarded as a very stable phenomenon. Just as any species has acquired a fixed and practically invariable size, so it would have acquired a definite longevity. 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 duration 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 100 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[29] 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 themselves. 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, 100 years; the camel takes 8 to grow, and lives 5 times as long, i.e., 40 years; the horse, 5 to grow, and lives 25 years.

However, even if we consider only the mammalia, it is impossible to accept Flourens’ law, without considerable reserve. Weismann[30] 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 reaching 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.[31] The longevity of the sheep, therefore, is not quite three times its period of growth.

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 reproduce, 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 quite remarkable longevity. 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 100 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.

H. Milne-Edwards[32] 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.”

Bunge[33] has recently taken up the study of the relations between the duration of growth and longevity, and has suggested a new means of investigation. 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. He has shown that whilst a human child requires 180 days to reach double its weight at birth, the horse, the longevity of which is very much less, doubles its weight in 60 days; a calf takes only 47 days for this; a kid 15 days; a pig 14 days; a cat 9-1/2; and a dog only 9 days. 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.

I observed myself that new-born mice quadruple their weight in the first 24 hours. 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 100 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. Now, it happens that there are some animals the fecundity of which is extremely limited. 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 circumstances the duration of life becomes a factor in the preservation 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 lead 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 100 years, or live beyond that time, much more often than men.

Low fertility, however, cannot itself be regarded as a cause of longevity, as there are some very fertile animals which none the less attain great ages. There are parrots which lay two or three times a year, producing six to nine eggs in each clutch. The ducks (Anatidæ) 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 carnivora. The longevity is possibly to be explained as an adaptation for the preservation of the species by compensating 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 independently. 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.

After having discussed various theories of the cause of the duration of life, M. Oustalet,[34] 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.” There are certainly many instances which give support to the view. Elephants and parrots, for instance, are vegetarian and reach very great ages. On the other hand, there exist long-living carnivorous 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.

II.
LONGEVITY IN THE ANIMAL KINGDOM

Longevity in the lower animals—Instances of long life in sea-anemones and other invertebrates—Duration of life of insects—Duration of life of “cold-blooded” vertebrates—Duration of life of birds—Duration of life of mammals—Inequality of the duration of life in males and females—Relations between longevity and fertility of the organism

It is wonderful to what an extent the duration of life varies amongst animals, the slightest examination of the facts showing that very many factors must be involved.

As the higher animals are nearly always larger than invertebrates, if there be a definite relation between longevity 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 was captured in 1828 by Dalyell, a Scottish zoologist, and was then quite adult, and probably about 7 years old. It survived its owner for 36 years, and died in Edinburgh in 1887, the cause of death being unknown. 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 1828 to 1848 produced 334 larvæ, then after a period of sterility it gave birth, in one night (1857) to 230 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 1872 onwards, it gave birth to 150 young anemones.[35] This animal, which certainly was not more than the fortieth or the fiftieth of the weight of an adult rabbit, lived six or seven times as long.

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.[36]

Insects are animals as variable in their duration of life as they are in other respects. Some live only a few weeks; some of the plant-lice, for instance, die in a month. 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 seventeen 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 grasshoppers 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.[37]

We know so little about the physiological processes of insects, that we cannot as yet make even a guess at the cause of this great variation in their longevity. It is more probable that we shall find some explanation in the case of vertebrates concerning which we know much more.

Analysis of the facts shows that whilst in the evolution from fish to mammal there has been a great increase in complexity of organisation, there has at the same time been a reduction in the duration of life. 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 numerous, 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 1230 and to have lived for 267 years afterwards. Carps are regarded as equally long lived, Buffon setting down their period of life as 150 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 estimate, inasmuch as during revolutionary times most of the carp were eaten when the palaces were overrun by the populace. 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. A tortoise has lived for eighty years in the garden of the Governor of Cape Town, and is believed to have reached the age of two hundred years. Another tortoise, a native of the Galapagos Islands, is known to be 175 years old, whilst a specimen in the London Zoological Gardens is 150 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 1623 and which lived there for 107 years.[38] Another tortoise, brought to Fulham by Archbishop Laud, lived in the Palace for 128 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 circulation, 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. J. H. Gurney.[39] In his list, in which are included more than fifty species of birds, the lowest figures are from eight and a half to nine years (Podargus cuvieri, Chelidon urbica), and a duration of life so short is an exception, a period of from fifteen to twenty years being more common. Canaries have lived in captivity 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 77. Jones, Layard, and Butler are the authorities for instances of Sulphur-crested Cockatoos having reached respectively 30, 72 and 81 years. M. Abrahams states that an Amazon (Chrysotis amasonica) lived 102 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 pneumonia.

Mr. Gurney found that parrots were not the only birds capable of reaching a great age. 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 common 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. Gurney. In the Royal Park at Schönbrunn, near Vienna, a white-headed vulture (Neophron percnopterus) died aged 118 years, a golden eagle (Aquila chrysaëtus) aged 104, and another aged 80 (according to Oustalet). Mr. Pycraft (Country Life, June 25th, 1904) reported that a female eagle, captured in Norway in 1829, 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 162 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 certainly 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 occasionally has completed its century. In zoological gardens and in good menageries, where elephants are well cared for, they seldom live more than 20 to 25 years. Chevrette, an African elephant presented to the Jardin des Plantes by Mehemet Ali, in 1825, lived for only 30 years. In the official list of the Indian Government, which gives the deaths of elephants, it appears that of 138 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 150 years to elephants as their epiphyses do not fuse with the long bones until the age of 30. So far, I know 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 140 years in which Ceylon was occupied by the Dutch. This elephant was found in the stables in 1656. Natives with special knowledge of elephants set down their duration of life as from 80 to 150 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. Grindon has stated his opinion that the rhinoceros may live for 70 or 80 years, but this seems rather an inference from the slowness of growth than a statement of observed fact.

Horses and cattle are large animals, but 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 46.

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 (242-287 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. Dogs seldom live more than 16 or 18 years, and even before that, at an age of from 10 to 12, they usually show plain signs of senility. Jonatt has mentioned as an extreme rarity a dog of 22 years of age, and Sir E. Ray Lankester (Comparative Longevity, p. 60) cites another instance, in this case the age being 34 years. The oldest dog that I have been able to procure died at the age of 22.

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. It is well known that an animal may be made feeble by too great fecundity, and it is conceivable that the kind of parasitic life of the embryos within the body of the mother may weaken her system.

There are many facts, however, which make it impossible 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 two sexes being specially striking in species of insects. Generally, the females live longer than the males, as, for instance, amongst the Strepsiptera, where the females have 64 times the duration of life of the males. On the other hand, amongst butterflies, there are cases (e.g., Aglia tau) where the males live longer than the females. 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 difference 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 reproductive 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. Fish or frogs which lay thousands of eggs at a time (a pike, for example, produces 130,000) are obviously more prolific than, for instance, a sparrow which lays only 18 eggs in a year, or than a rabbit, which in the same time gives birth to from 25 to 50. 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.

III
THE DIGESTIVE SYSTEM AND SENILITY

Relations between longevity and the structure of the digestive system—The Cæca in birds—The large intestine of mammals—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

We have seen that the duration of life in mammals is relatively shorter than that in birds, and in the so-called “cold-blooded” vertebrates. 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 intestine 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 cæcum. In birds, the large intestine still remains relatively badly developed; it is short and straight. In most birds, at the point where the large intestine passes into the small intestine, there is a pair of cæca, more or less developed. These cæca are absent in climbing birds, such as the wood-pecker, the oriole, and many others. They are reduced to a pair of tiny outgrowths 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.[40]

In the large running birds, such as ostriches, rheas, and tinamous, the cæca are relatively largest. Thus, for instance, in a rhea (Rhea americana) which I dissected, the cæca were nearly two-thirds as long as the small intestine. The latter was 1·65 m. in length, whereas one of the cæca was 1·01 m., and the other 0·95 m. The weight of the two cæca 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.”[41]

Gegenbaur,[42] another well-known authority on comparative anatomy, writes as follows on this subject:—“The hind-gut is longest in the Mammalia, where it forms the large intestine, and is distinguished as such, from the mid-gut, or small intestine. Owing to 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.”

The two series of facts are not to be disputed. 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.), the large intestine is not more than a mere reservoir for the waste matter in the food. It takes no share in digestion, as that is the function of the stomach and the small intestine. Only the cæcum can be thought to have some digestive property. In reptiles, the lowest vertebrates in which the cæcum 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, however, the cæca are well separated from the main digestive tube. The food material passes into them in considerable quantities, and is retained there sufficiently long for some digestive process to take place. M. Maumus has found, in the cæca of birds, secretions which can dissolve albumen and invert sugar cane, but he has been unable to make out that the cæcal juice has any action upon fatty matter. Such digestive power, however, is slight, and when M. Maumus removed the cæca in fowls and ducks, no evil consequences followed. As in many birds the cæca are rudimentary and in others absent, it may be inferred that these organs are useless, and are in process of degeneration in the class. The cæca can be regarded as playing an important part in the organism only in the case of large running birds, where they are very highly developed, but we have not precise information as to their digestive function.

The variations in the structure in the large intestine are greater in mammals than in birds. In some mammals, the large intestine is a simple prolongation of the small intestine, similar in calibre and in structure. In these conditions it may fulfil a definite digestive function. Th. Eimer[43] has determined that in insectivorous bats the large intestine digests insects like the small intestine. Such cases, however, are rare. In most mammals the large intestine is sharply separated from the small intestine by a valve, and opens directly into the cæcum which may be very large. In the horse, the cæcum is an enormous bag, cylindrical and tapering, generally well filled, and holding on an average 35 litres. It is equally large in many other herbivorous animals, such as the tapir, the elephant, and most rodents. In such cases, the food remains for a considerable time in the organ and without doubt undergoes some digestive changes. In many other mammals, particularly carnivorous forms, the cæcum may be quite absent, whilst in some, as for instance, the cat and dog, it is very small; in the latter cases its digestive function must be non-existent or insignificant.[44]

As for the large intestine itself, apart from the special cases, such as bats, it cannot fulfil any notable digestive function. Th. Eimer was unable to find a proof of any such action in rats and mice, and the very many investigations that have been made in the case of man seem to have established the absence of digestive power in the colon.

Dr. Stragesco,[45] in a recent investigation carried out under the direction of the famous Russian physiologist Pawloff, established that, in normal conditions, digestion and assimilation of food are confined almost exclusively to the small intestine in mammals, and that the large intestine plays only the smallest part. It is only in certain diseases of the digestive tract, in which, on account of increased peristaltic action, the contents of the intestine with the digestive juices are passed quickly from the small intestine to the large intestine, that some digestive work is done in the latter organ.

The large intestine (excluding the cæcum), then, cannot be regarded as an organ of digestion, although absorption of the liquids which have been formed in the small intestine, may take place within its walls. It is known that in the large intestine the contents of the gut give up their water and assume the solid form of fæcal matter. However, whilst the mucous membrane of the large intestine rapidly absorbs water, it has not a similar action on other substances.

The question of the extent to which the large intestine can absorb has been closely investigated, because of its practical importance. It sometimes happens that invalids cannot take food by the mouth, so that their life would be in danger if it were not possible to supply them with food otherwise. Attempts have been made to inject nutritive substances through the skin, or, and this is a more usual procedure, by the rectum. By such means the organism can be kept alive for a certain time, but the absorbing power of the large intestine is extremely small. According to Czerny and Lautschenberger[46] the entire colon of the human being can absorb no more than 6 grammes of albumen in 24 hours, an amount which, from the point of view of nutrition, is very small. It was thought that the large intestine might more rapidly absorb albuminous material which had been previously digested and transformed to peptones, but the experiments of Ewald[47] showed that even in that case the absorption was very small. According to more recent experiments of Heile,[48] carried out upon dogs which had cæcal fistulas, and in the case of a man who had an artificial aperture in the colon, the large intestine does not absorb undigested white of egg, and absorbs water, cane sugar, and glucose only very imperfectly. The only substances which are rapidly absorbed through the wall of the colon are the alkaline fluids from fæcal matter. It is possible, however, to nourish invalids by rectal injections of certain nutritious substances, the most important of which is milk.[49]

The large intestine, which has really very slight digestive properties and cannot absorb any considerable bulk of nutriment, is an organ which secretes mucus. The latter serves to moisten the solid fæcal material, so aiding in its expulsion.

We must conclude, therefore, that the large intestine, the organ so highly developed in mammals, is an apparatus the general function of which is the preparation and elimination of the waste products of digestion. Why should such an organ be so much more developed in mammals than in the other vertebrates?

In answer to the question, I have formed the theory that the large intestine has been increased in mammals to make it possible for these animals to run long distances without having to stand still for defæcation. The organ, then, would simply have the function of a reservoir of waste matter.

Batrachia and reptiles lead a very idle life, and can move slowly, sometimes because they are protected by poison (toads, salamanders, serpents), sometimes because they have a very hard shell (turtles), sometimes because they are extremely powerful (crocodiles). Mammals, on the other hand, have to move very actively to catch their prey, or to escape from their enemies. Such activity has become possible because of the high development of the limbs, and because the capacity of the large intestine makes possible the accumulation of waste matter for a considerable time.

In order to void the contents of the intestines, mammals have to stand still and assume some particular position. Each act of this kind is a definite risk in the struggle for existence. A carnivorous mammal which, in the process of hunting its prey, had to stop from time to time, would be inferior to one which could pursue its course without pausing. So, also, a herbivorous mammal, escaping from an enemy by flight, would have the better chance of surviving the less it was necessary for it to stand still.

According to such a view, the extreme development of the large intestine would supply a real want in the struggle for existence. M. Yves Delage,[50] the well-known biologist, is unable to accept this hypothesis. He thinks that the rectal enlargement would fulfil the purpose, and adds that everyone has seen herbivorous animals pass their excretions whilst running. The rectum of mammals, however, cannot serve as a reservoir for waste matter, because as soon as such matter reaches the rectum it excites the need of excretion. The waste matter accumulates in the large intestine, from which it passes into the rectum at intervals. When it has reached that region, a sensation is caused which leads to defæcation.

M. Delage is not quite definite when he speaks of mammals voiding their excretions whilst they are in motion. A horse, harnessed to a vehicle, may defæcate whilst it is walking or even running slowly. But these animals cannot defæcate when in rapid motion, and competent observers state that horses never do so whilst racing. In zoological gardens, where animals have room to run about, they stand still before emptying the rectum. M. Ch. Debreuil, who keeps antelopes in a very large park at Melun, has noticed that the excreta are always to be found in masses and not scattered about as if they had been discharged by animals in motion. Antelopes, which are animals that run and leap extremely actively, have to come to a standstill before discharging their small pellets of deer-like excreta.

In the struggle for existence, when a mammal is pursuing its prey or escaping its enemy, there is no question of the leisurely movement of a horse harnessed to an omnibus or cab, but the greatest possible activity is necessary. In such circumstances the possession of an organ within which the excreta could accumulate would be of real importance. My theory of the origin of the mammalian large intestine is intrinsically probable.

Although the capacity of the large intestine may preserve a mammal in emergencies, it is attended with disadvantages that may shorten the actual duration of life.

The accumulation of waste matter, retained in the large intestine for considerable periods, becomes a nidus for microbes which produce fermentations and putrefaction harmful to the organism. Although our knowledge of the subject is far from complete, it is certain that the intestinal flora contains some microbes which damage health, either by multiplying in the organism, or by poisoning it with their secretions. Most of our knowledge on this matter has come from the study of human patients.

Persons have been known who do not defæcate except at intervals of several days, and who, none the less, do not seem to suffer in health. But the opposite result is more common. The retention of fæcal matter for several days very often brings harmful consequences. Organisms which are in a feeble state from some other cause are specially susceptible to damage of the kind referred to. Infants are frequently seriously ill as the result of constipation. Dr. du Pasquier[51] describes such cases in the following words:—“The infant is leaden in hue, with sunken eyes, dilated pupils, and pinched nostrils. The temperature may reach nearly 104° Fahr.; the pulse is rapid, feeble, and often irregular. Restlessness, insomnia, sometimes convulsions, stiffness of the neck and strabism show that the nervous system is being poisoned by toxins, and even collapse may be reached. The foul and dry tongue, the vomiting and fetid discharges show the disturbance of the digestive tract. Very often an eruption appears, as described by Hutinel, chiefly on the back and buttocks, the front of the thighs and fore-arms.” The illness may lead to death but is generally cured by simple purging.

Women in pregnancy and child-birth frequently suffer much as the result of retention of fæcal matter, and physicians are familiar with the symptoms, which have been described as follows by M. Bouchet[52]:—“After normal parturition, in the course of which the usual antiseptic precautions have been fully pursued, and where delivery has been complete and natural, occasionally the patient is seized with chill and headache. The breath is fetid and the tongue foul. The temperature, taken in the axilla, is nearly 101° Fahr. The abdomen is inflated and painful in the umbilical region. Palpation in the iliac fossæ reveals lumps or consolidations along the colon. Thirst is intense, and there is complete anorexy. On questioning, it is found that there has not been defæcation for several days. The treatment consists of purgatives, enemas, and milk diet. In the next few days the bowels are emptied freely, the abdominal pain ceases, the temperature becomes lower, appetite is restored, and the patient recovers.”

Those who suffer from affections of the heart, liver, or kidneys are specially susceptible to the evil results of retained fæcal matter. In such patients an error of diet or constipation may bring about most serious consequences.

Such facts are well known to physicians, and it has been established that complete emptying of the lower bowels leads at once to favourable symptoms. From the other side, it has been shown by experiment that artificial retention of the fæces by ligature of the rectum puts the body in a grave condition.

If we collect our knowledge of all the facts, we cannot doubt but that the cause of the evil is multiplication of microbes in the contents of the large intestine. When the fæcal matter is free from microbes, as is the case with the meconium of the fœtus or new-born infant, it is not a source of danger to the organism. The waste of cells and the secretions which are added to the undigested food cannot do any harm. Amongst the microbes of the gut, there are some that are inoffensive, but others are known to have pernicious properties.

The ill-health which follows retention of fæcal matter is certainly due to the action of some of the microbes of the gut. There are difficulties, however, in determining the precise mode of action of these microbes. It is generally believed that they form poisonous substances which are absorbed by the walls of the intestine and so pass into the system. The phrase auto-intoxication as applied to infants, women in labour, and patients affected with diseases of the heart, liver, or kidneys, is based on this interpretation of the morbid processes involved. Attempts have been made to isolate and study the poisons in question, but there are many difficulties in the way. To distinguish between the actions of the poisons and of the microbes themselves, the latter have been destroyed by heat or by antiseptics, or been removed by filtration. Such methods, however, may alter the poisons and so are inconclusive. MM. Charron and Le Play[53] have tried to obtain exact results by heating the intestinal microbes to a temperature of about 136° Fahr., a process which probably does not seriously deteriorate the microbial poisons. Such material, injected into the veins of rabbits in large quantities, rapidly produced death, or in smaller quantities, proportionate ill-health.

Kukula[54] has tried to produce this toxic action in animals, employing microbial secretions obtained from cases of intestinal obstruction. He succeeded in producing serious symptoms, such as vomiting and curvature of the neck and back, in fact, precisely the sequence of events familiar in cases of obstruction of the bowels or other retentions of fæcal matter.

Some of the products of the intestinal flora are undoubtedly toxic, such as the benzol derivatives (phenol, etc.) ammonium and other salts. Many of these toxins have been insufficiently studied, but it is well known that certain of them can be absorbed by the wall of the gut and act as poisons. A well known case is the toxin of botulism which was isolated and studied by M. van Ermenghem.[55] The poison, the product of a microbe which causes serious intestinal disturbance, is so fatal that a single drop given to a rabbit produces death after symptoms similar to those observed in cases of human beings poisoned by stale food. Butyric acid and the products of albuminous putrefaction are amongst the most pernicious of the microbial poisons produced in the large intestine. It is familiar that digestive disturbance is frequently associated with discharges of sulphuretted hydrogen and putrid excreta, and there is no doubt but that the microbes of putrefaction are the cause of these symptoms.

It has been assumed for long that the retention of fæcal matter tends to putrefactive changes in the intestines, and that the evil consequences of constipation are due to this. Recently, however, bacteriologists have criticised this accepted view, on account of the small number of microbes found in the excreta of constipated persons. Strasburger was the first to establish the fact, and his associate, Schmidt, showed that putrefaction did not follow when readily putrescible substances were infected with material taken from cases of constipation. However, notwithstanding the exactness of these facts, I cannot accept the inference which has been drawn from them. The excreta discharged naturally in cases of constipation do not give a correct indication of the conditions inside the gut; whilst such matter contains few microbes, the substance removed after injection by an enema is extremely rich in bacteria. Moreover, analysis of the urine, in cases of constipation, shows an excess of the sulpho-conjugate ethers which are known to be products of intestinal putrefaction.

Not only is there auto-intoxication from the microbial poisons absorbed in cases of constipation, but microbes themselves may pass through the walls of the intestine and enter the blood. In the maladies that are the result of constipation some of the symptoms recall those of direct infection, and it is highly probable that, if special investigations were made, microbes of intestinal origin would be found in the blood of the sick children and the pregnant or parturient women whose symptoms I have described above.

The question as to the passage of microbes through the intestinal walls is one of the most controversial of bacteriological problems, and there is little agreement in the numerous publications regarding it. None the less, it is far from impossible to get a general idea of what goes on in an intestinal tract richly charged with microbes.

Although the intestinal wall in an intact state offers a substantial obstacle to the passage of bacteria, it is incontestable that some of these pass through it into the organs and the blood. Numerous experiments performed on different kinds of animals (horses, dogs, rabbits, etc.) show that some of the microbes taken with food traverse the wall of the alimentary canal and come to occupy the adjacent lymphatic glands, the lungs, the spleen and the liver, whilst they are occasionally found in the blood and lymph. Discussion has taken place as to whether the passage takes place when the wall of the gut is absolutely intact or only when it is injured to however small an extent. It would be extremely difficult to settle the question definitely, but it is easy to see that it has little practical bearing. It is known that the wall of the gut is damaged extremely easily, so that the bluntest sound can hardly be passed into the stomach without making a wound through which microbes can pass into the tissues and blood. In the ordinary course of life, the delicate wall of the gut must often undergo slight wounding, and the frequent presence of microbes in the mesenteric ganglia of healthy animals shows clearly what takes place.[56]

It is indubitable, therefore, that the intestinal microbes or their poisons may reach the system generally and bring harm to it. I infer from the facts that the more a digestive tract is charged with microbes, the more it is a source of harm capable of shortening life.

As the large intestine not only is the part of the digestive tube most richly charged with microbes, but is relatively more capacious in mammals than in any other vertebrates, it is a just inference that the duration of life of mammals has been notably shortened as the result of chronic poisoning from an abundant intestinal flora.

IV
MICROBES AS THE CAUSE OF SENILITY

Relations between longevity and the intestinal flora—Ruminants—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

In the actual state of our knowledge it is impossible to make a final examination of my hypothesis, as there are many factors about which we are incompletely informed. Nevertheless, it is possible to confront the hypothesis with a large number of accurately established facts.

Although the life of most mammals is relatively short, there are to be found in the group some which live relatively long, as well as others whose life is short. The elephant is an example of the long-lived mammals, whilst ruminants are short-lived forms. In the last chapter, I stated that sheep and cattle became senile at an early age, and did not live long. They are striking exceptions to the rule according to which the duration of life is in direct relation with the size and length of the period of growth. The cow, which is much larger than a woman, and the time of gestation of which is about the same, or a little longer, acquires its teeth at four years old, and becomes senile at an early age; it is quite old at between sixteen and seventeen, an age when a woman is hardly adult; at the age of thirty, practically the extreme limit for bovine animals, a woman is in full vigour.

The precocious old age of ruminants, the constitution of which is well understood, and which are carefully tended, coincides with an extraordinary richness of the intestinal flora. Food remains for a long time in the complicated stomach of these animals, and afterwards the digested masses remain still longer in the large intestine. According to Stohmann and Weiske,[57] in the case of sheep it is a week until the remains of a particular meal have finally left the body of the animal. The excreta of sheep, normally solid, do not betray any special putrefaction in the intestine, but if the body is opened there is abundant evidence of the process. The intestinal contents are richly charged with microbes and give off a strong odour of putrefaction. It is not surprising that under these conditions, the life of sheep should be short.

Another large herbivorous animal, the horse, also dies young, after a premature old age. Although it does not ruminate and possesses a simple stomach, the process of digestion is slow, and enormous masses of nutritive material accumulate in the huge large intestine. Ellenberger and Hofmeister[58] have shown that food remains in the alimentary canal for nearly four days. It remains in the stomach and the small intestine only 24 hours, but about three times as long in the large intestine. This is remarkably different from what happens in the case of birds, in which there is no stagnation during the passage of food through the digestive canal.

The structure of birds is adapted for flight, the body being as light as possible, many of the bones and the cavities of the body containing air-sacs. The absence of a bladder and of a true large intestine prevents the accumulation of excreta, these being ejected almost as rapidly as they are formed. The process of ejection, which takes place often in birds, is not so inconvenient as in mammals. The hind limbs are not used in flight, so that they offer no obstacle to evacuation. Thus birds may discharge their droppings while flying.

Such structure and habits make it not surprising that the alimentary canal of many birds contains only a scanty intestinal flora. Parrots, for instance, which are remarkably long-lived birds, harbour very few microbes in the intestine. The small intestine contains almost none, the rectum so few that the fæcal matter appears to be formed of mucus, the waste of the food, and only a very few microbes. M. Michel Cohendy, who has examined the intestinal flora at the Pasteur Institute, was unable to isolate more than five different species of microbes living in the alimentary canal of parrots.

Even in birds of prey which feed upon putrid flesh, the number of microbes in the intestine is remarkably limited. I have investigated the case of ravens which I fed on flesh which was putrid and swarming with microbes. The droppings contained very few bacteria, and it was specially remarkable that the intestines had not the slightest smell of putrefaction. Although the opened body of a herbivorous mammal, such as a rabbit, gives off a strong smell of putrefaction, the body of a raven with the digestive tube exposed has no unpleasant smell. This absence of putrefaction in the intestine is probably the reason of the great longevity of such birds as parrots, ravens, and their allies.

It might be said, however, that the long duration of life in birds is due to the organisation of these animals, rather than to the scantiness of their intestinal flora. To meet this objection, it is necessary to turn to the case of cursorial birds.

There are some birds incapable of flight, the wings of which are badly developed, but which have strong limbs, and can run with great rapidity. Ostriches, cassowaries, rheas, and tinamous, are well known examples of cursorial birds. They live on the surface of the ground, and their habits resemble those of mammals. When they are attacked by enemies, they escape by running so quickly that some of them (ostriches and rheas) outstrip even a horse. However, like mammals, they cannot discharge their secretions when they are running quickly. Tinamous (Rhynchotus rufescens), which I have observed in captivity, however quickly they may be running, stop abruptly to discharge their excretions. M. Debreuil, at my request, made observations on this matter, and assured me that the tinamous and rheas (Rhea americana) in his park always stood still for this purpose. He has noticed that the droppings, however abundant, were always deposited in heaps. With regard to ostriches, M. Rivière, director of the experimental Gardens at Hamma, Algeria, has been kind enough to give me the following information. “The discharge of excreta,” he said in a letter in January, 1901, “is less frequent than in other birds, but the comparatively small size of the enclosures here makes it impossible for me to assert that the animal could discharge its droppings if it were running for a length of time; a priori I should think that this did not happen. Normally the bird stands still for defæcation, the tuft of feathers on the tail is lifted up, and there is a violent contraction of the abdominal muscles before the sphincters of the cloaca are suddenly opened to discharge the excrement with violence.”

I believe that the remarkable development of the large intestine in these running birds has been acquired to obviate the danger which is caused by the animal having to stop for defæcation. Although the huge cæca of these birds have a digestive function, particularly on plants rich in cellulose, I cannot think that the cæca of cursorial birds have been developed for digestion. As a matter of fact, some birds which are not cursorial live on the same kind of food (herbage, seeds, and insects) and have much smaller cæca, the cæca indeed, in some, for instance, the pigeons, being quite rudimentary.

Fig. 14.—Intestinal microbes from the cæca of a Rhea.

It is not surprising that the accumulation of food material in the large intestine of running birds is associated with the presence of an extremely rich intestinal flora. Microscopic examination of the excrement of such birds shows this at once. Although the intestinal contents and excrement of many other birds show the presence of very few microbes, belonging to a small number of species, the same materials taken from running birds show enormous quantities of microbes, belonging to a large number of species. In the cæcum of the rhea (Fig. [14]) there are bacterial threads, spirilla, bacilli, vibrios, and many kinds of cocci. In the tinamous, the intestinal flora is if possible even richer. According to the statistical investigations of M. Michel Cohendy, the quantity of intestinal microbes in cursorial birds is not less than that found in mammals, even in man.

If I am correct in the view that I have been explaining, cursorial birds, on account of their rich intestinal flora, ought to have a shorter duration of life than that of flying birds. I will now turn to this side of the question. Amongst cursorial forms, there are some of the largest living birds, ostriches being actually the largest living birds, whilst an extinct running bird, the Aepyornis of Madagascar, was the largest known bird. According to the rule that large animals live longer than small animals, ostriches should be able to reach a great age. The facts, however, are against this. M. Rivière, who rears ostriches in Algeria, and has a great experience of them, writes to me as follows: “I have no confidence in the stories about the longevity of the ostrich which were told me in the Sahara; they rest on no facts. My personal observation is not very large, but it is quite exact. Some of the ostriches which have been hatched here have lived for 26 years. I do not estimate the duration of life of this bird at more than 35 years, and only one case of this age have I seen myself in 20 years. The bird was a female, a good layer and sitter; she died of old age, showing all the signs of decrepitude, the skin excoriated and lumpy, the feathers degenerate and dry. The bird laid eggs until nearly the end of her life, but at irregular intervals, and the shells were granular instead of being smooth and polished.”

In a farm near Nice, where ostriches are reared, there was recently an old male called “Kruger,” which was supposed to be 50 years old.[59] Countess Stackelberg has been good enough to try to get information for me about this, and informs me that although they have not exact knowledge at the farm, they believe that it must be 50 years old. M. Rivière thinks this statement very surprising, and has nothing in his own long experience to confirm it.

The facts which I have been able to get together do not attribute a long life to other running birds. Gurney mentions that a cassowary (Casuarius westermanni) lived 26 years in the Zoological Gardens of Rotterdam, and that three Australian emus (Dromaeus novae-hollandiae) had lived in the same Gardens for 28, 22, and 20 years. M. Oustalet (Ornis, 1899, vol. x, p. 62) mentions another emu of the same species which died in London at the age of over 23 years. The rhea (Rhea americana), another large running bird, does not live so long. “Boecking thinks that its duration of life should be set down at from 14 to 15 years. According to him, many of these birds die of old age.” (Brehm, Oiseaux, vol. ii, p. 517).

It is striking to compare the short life of cursorial birds, which nevertheless thrive and reproduce in captivity, with the remarkable longevity of so many other birds (parrots, birds of prey) which, although they are much smaller, have been kept alive for from 80 to 100 years. It would be difficult to find a more striking argument in favour of the view that richness of the intestinal flora shortens life. When birds become adapted to terrestrial life and acquire a huge large intestine in which microbes can abound, their duration of life is diminished.

Just as some birds, losing the aerial mode of life, have come to resemble mammals, so also some mammals have become flying animals, provided with wings and in some respects resembling birds. Bats are the most familiar instance. The large intestine, which is extremely useful to running animals, not only ceases to be an advantage but is harmful to flying creatures, insomuch as it increases the weight of the body uselessly. Bats, accordingly, have no cæcum whilst the large intestine is changed in structure and function. Instead of being a capacious tube, serving as a reservoir for the refuse of the food, the large intestine of bats has the same diameter as the small intestine. Its structure is nearly identical. It is provided with glands, and as I have already mentioned in the last chapter, it digests the food in the same way as the small intestine. In fact, the large intestine has become simply a part of the small intestine, the total length of the gut being reduced. Bats, therefore, can no longer retain their secretions but have to empty the intestine almost as often as most birds. I find that Indian fruit bats (Pteropus medius) discharge their excreta very often. Microscopic examination shows that there is an absence of microbes quite unusual in the case of a mammal. The alimentary canal of bats is nearly aseptic, containing only a few single bacteria. I have fed these fruit bats with the same food (carrots) which I have given to rabbits, guinea pigs, and mice; whilst the bats accomplished the process of digestion in 1-1/2 hours, and deposited excreta containing fragments of carrot, the rodents took very much longer for digestion and large quantities of waste matter accumulated in the cæca. The intestinal flora too, although the food in each case was the same, showed remarkable differences in these animals. It was almost absent in the bats, whilst in the rabbits, guinea-pigs and mice it consisted of a mass of microbes of different species. The excrement of the bats had no unpleasant odour, and the digestive canal of these bird-like mammals was free from putrefaction. Fruit bats fed upon fruit discharged excreta with a pleasant odour of apples and bananas. We have seen that birds which live a life similar to that of mammals acquire a rich intestinal flora and do not live so long as aerial birds. It would be extremely interesting to ascertain the duration of life of bats, mammals which live like birds and have a very scanty intestinal flora. I have been unable to get any exact information as to the duration of life of the true bats, that is to say, the insectivorous bats, as all the requests that I have addressed to specialists have proved fruitless. It appears, however, that it is a popular belief that bats live long. There is a Flemish phrase: “as long-lived as a bat,” and a similar phrase is common in Little Russia.

As for the fruit-eating bats, I have been able to ascertain that even in captivity, where the conditions are unfavourable to them, the duration of life is relatively long. I have had in my own possession a fruit bat (Pteropus medius) which was bought in Marseilles 14 years ago. It showed no signs of old age, and the teeth were in perfect condition. It died of some acute disease accidentally contracted. I know of another bat of the same species which lived in captivity for more than 15 years, and I have been informed that[60] in the London Zoological Gardens, a fruit bat has lived for 17 years. If these bats were adult when caught, it would be necessary to add something to the known figures.

Although I do not know the exact duration of the life of bats, it is clearly relatively long for mammals no bigger than guinea-pigs. The difference is remarkable if we compare it with the life of sheep, dogs and rabbits, mammals very much larger in size, but possessed of a rich intestinal flora.