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  • 1904
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many years of experimentation brought it to relative perfection in 1839, in which year the famous daguerreotype first brought the matter to popular attention. In the same year Mr. Fox Talbot read a paper on the subject before the Royal Society, and soon afterwards the efforts of Herschel and numerous other natural philosophers contributed to the advancement of the new method.

In 1843 Dr. John W. Draper, the famous English-American chemist and physiologist, showed that by photography the Fraunhofer lines in the solar spectrum might be mapped with absolute accuracy; also proving that the silvered film revealed many lines invisible to the unaided eye. The value of this method of observation was recognized at once, and, as soon as the spectroscope was perfected, the photographic method, in conjunction with its use, became invaluable to the chemist. By this means comparisons of spectra may be made with a degree of accuracy not otherwise obtainable; and, in case of the stars, whole clusters of spectra may be placed on record at a single observation.

As the examination of the sun and stars proceeded, chemists were amazed or delighted, according to their various preconceptions, to witness the proof that many familiar terrestrial elements are to be found in the celestial bodies. But what perhaps surprised them most was to observe the enormous preponderance in the sidereal bodies of the element hydrogen. Not only are there vast quantities of this element in the sun’s atmosphere, but some other suns appeared to show hydrogen lines almost exclusively in their spectra. Presently it appeared that the stars of which this is true are those white stars, such as Sirius, which had been conjectured to be the hottest; whereas stars that are only red-hot, like our sun, show also the vapors of many other elements, including iron and other metals.

In 1878 Professor J. Norman Lockyer, in a paper before the Royal Society, called attention to the possible significance of this series of observations. He urged that the fact of the sun showing fewer elements than are observed here on the cool earth, while stars much hotter than the sun show chiefly one element, and that one hydrogen, the lightest of known elements, seemed to give color to the possibility that our alleged elements are really compounds, which at the temperature of the hottest stars may be decomposed into hydrogen, the latter “element” itself being also doubtless a compound, which might be resolved under yet more trying conditions.

Here, then, was what might be termed direct experimental evidence for the hypothesis of Prout. Unfortunately, however, it is evidence of a kind which only a few experts are competent to discuss–so very delicate a matter is the spectral analysis of the stars. What is still more unfortunate, the experts do not agree among themselves as to the validity of Professor Lockyer’s conclusions. Some, like Professor Crookes, have accepted them with acclaim, hailing Lockyer as “the Darwin of the inorganic world,” while others have sought a different explanation of the facts he brings forward. As yet it cannot be said that the controversy has been brought to final settlement. Still, it is hardly to be doubted that now, since the periodic law has seemed to join hands with the spectroscope, a belief in the compound nature of the so-called elements is rapidly gaining ground among chemists. More and more general becomes the belief that the Daltonian atom is really a compound radical, and that back of the seeming diversity of the alleged elements is a single form of primordial matter. Indeed, in very recent months, direct experimental evidence for this view has at last come to hand, through the study of radio-active substances. In a later chapter we shall have occasion to inquire how this came about.



An epoch in physiology was made in the eighteenth century by the genius and efforts of Albrecht von Haller (1708-1777), of Berne, who is perhaps as worthy of the title “The Great” as any philosopher who has been so christened by his contemporaries since the time of Hippocrates. Celebrated as a physician, he was proficient in various fields, being equally famed in his own time as poet, botanist, and statesman, and dividing his attention between art and science.

As a child Haller was so sickly that he was unable to amuse himself with the sports and games common to boys of his age, and so passed most of his time poring over books. When ten years of age he began writing poems in Latin and German, and at fifteen entered the University of Tubingen. At seventeen he wrote learned articles in opposition to certain accepted doctrines, and at nineteen he received his degree of doctor. Soon after this he visited England, where his zeal in dissecting brought him under suspicion of grave-robbery, which suspicion made it expedient for him to return to the Continent. After studying botany in Basel for some time he made an extended botanical journey through Switzerland, finally settling in his native city, Berne, as a practising physician. During this time he did not neglect either poetry or botany, publishing anonymously a collection of poems.

In 1736 he was called to Gottingen as professor of anatomy, surgery, chemistry, and botany. During his labors in the university he never neglected his literary work, sometimes living and sleeping for days and nights together in his library, eating his meals while delving in his books, and sleeping only when actually compelled to do so by fatigue. During all this time he was in correspondence with savants from all over the world, and it is said of him that he never left a letter of any kind unanswered.

Haller’s greatest contribution to medical science was his famous doctrine of irritability, which has given him the name of “father of modern nervous physiology,” just as Harvey is called “the father of the modern physiology of the blood.” It has been said of this famous doctrine of irritability that “it moved all the minds of the century–and not in the departments of medicine alone–in a way of which we of the present day have no satisfactory conception, unless we compare it with our modern Darwinism.”[1]

The principle of general irritability had been laid down by Francis Glisson (1597-1677) from deductive studies, but Haller proved by experiments along the line of inductive methods that this irritability was not common to all “fibre as well as to the fluids of the body,” but something entirely special, and peculiar only to muscular substance. He distinguished between irritability of muscles and sensibility of nerves. In 1747 he gave as the three forces that produce muscular movements: elasticity, or “dead nervous force”; irritability, or “innate nervous force”; and nervous force in itself. And in 1752 he described one hundred and ninety experiments for determining what parts of the body possess “irritability”–that is, the property of contracting when stimulated. His conclusion that this irritability exists in muscular substance alone and is quite independent of the nerves proceeding to it aroused a controversy that was never definitely settled until late in the nineteenth century, when Haller’s theory was found to be entirely correct.

It was in pursuit of experiments to establish his theory of irritability that Haller made his chief discoveries in embryology and development. He proved that in the process of incubation of the egg the first trace of the heart of the chick shows itself in the thirty-eighth hour, and that the first trace of red blood showed in the forty-first hour. By his investigations upon the lower animals he attempted to confirm the theory that since the creation of genus every individual is derived from a preceding individual–the existing theory of preformation, in which he believed, and which taught that “every individual is fully and completely preformed in the germ, simply growing from microscopic to visible proportions, without developing any new parts.”

In physiology, besides his studies of the nervous system, Haller studied the mechanism of respiration, refuting the teachings of Hamberger (1697-1755), who maintained that the lungs contract independently. Haller, however, in common with his contemporaries, failed utterly to understand the true function of the lungs. The great physiologist’s influence upon practical medicine, while most profound, was largely indirect. He was a theoretical rather than a practical physician, yet he is credited with being the first physician to use the watch in counting the pulse.


A great contemporary of Haller was Giovanni Battista Morgagni (1682-1771), who pursued what Sydenham had neglected, the investigation in anatomy, thus supplying a necessary counterpart to the great Englishman’s work. Morgagni’s investigations were directed chiefly to the study of morbid anatomy–the study of the structure of diseased tissue, both during life and post mortem, in contrast to the normal anatomical structures. This work cannot be said to have originated with him; for as early as 1679 Bonnet had made similar, although less extensive, studies; and later many investigators, such as Lancisi and Haller, had made post-mortem studies. But Morgagni’s De sedibus et causis morborum per anatomen indagatis was the largest, most accurate, and best-illustrated collection of cases that had ever been brought together, and marks an epoch in medical science. From the time of the publication of Morgagni’s researches, morbid anatomy became a recognized branch of the medical science, and the effect of the impetus thus given it has been steadily increasing since that time.


William Hunter (1718-1783) must always be remembered as one of the greatest physicians and anatomists of the eighteenth century, and particularly as the first great teacher of anatomy in England; but his fame has been somewhat overshadowed by that of his younger brother John.

Hunter had been intended and educated for the Church, but on the advice of the surgeon William Cullen he turned his attention to the study of medicine. His first attempt at teaching was in 1746, when he delivered a series of lectures on surgery for the Society of Naval Practitioners. These lectures proved so interesting and instructive that he was at once invited to give others, and his reputation as a lecturer was soon established. He was a natural orator and story-teller, and he combined with these attractive qualities that of thoroughness and clearness in demonstrations, and although his lectures were two hours long he made them so full of interest that his pupils seldom tired of listening. He believed that he could do greater good to the world by “publicly teaching his art than by practising it,” and even during the last few days of his life, when he was so weak that his friends remonstrated against it, he continued his teaching, fainting from exhaustion at the end of his last lecture, which preceded his death by only a few days.

For many years it was Hunter’s ambition to establish a museum where the study of anatomy, surgery, and medicine might be advanced, and in 1765 he asked for a grant of a plot of ground for this purpose, offering to spend seven thousand pounds on its, erection besides endowing it with a professorship of anatomy. Not being able to obtain this grant, however, he built a house, in which were lecture and dissecting rooms, and his museum. In this museum were anatomical preparations, coins, minerals, and natural-history specimens.

Hunter’s weakness was his love of controversy and his resentment of contradiction. This brought him into strained relations with many of the leading physicians of his time, notably his own brother John, who himself was probably not entirely free from blame in the matter. Hunter is said to have excused his own irritability on the grounds that being an anatomist, and accustomed to “the passive submission of dead bodies,” contradictions became the more unbearable. Many of the physiological researches begun by him were carried on and perfected by his more famous brother, particularly his investigations of the capillaries, but he added much to the anatomical knowledge of several structures of the body, notably as to the structure of cartilages and joints.


In Abbot Islip’s chapel in Westminster Abbey, close to the resting-place of Ben Jonson, rest the remains of John Hunter (1728-1793), famous in the annals of medicine as among the greatest physiologists and surgeons that the world has ever produced: a man whose discoveries and inventions are counted by scores, and whose field of research was only limited by the outermost boundaries of eighteenth-century science, although his efforts were directed chiefly along the lines of his profession.

Until about twenty years of age young Hunter had shown little aptitude for study, being unusually fond of out-door sports and amusements; but about that time, realizing that some occupation must be selected, he asked permission of his brother William to attempt some dissections in his anatomical school in London. To the surprise of his brother he made this dissection unusually well; and being given a second, he acquitted himself with such skill that his brother at once predicted that he would become a great anatomist. Up to this time he had had no training of any kind to prepare him for his professional career, and knew little of Greek or Latin–languages entirely unnecessary for him, as he proved in all of his life work. Ottley tells the story that, when twitted with this lack of knowledge of the “dead languages” in after life, he said of his opponent, “I could teach him that on the dead body which he never knew in any language, dead or living.”

By his second year in dissection he had become so skilful that he was given charge of some of the classes in his brother’s school; in 1754 he became a surgeon’s pupil in St. George’s Hospital, and two years later house-surgeon. Having by overwork brought on symptoms that seemed to threaten consumption, he accepted the position of staff-surgeon to an expedition to Belleisle in 1760, and two years later was serving with the English army at Portugal. During all this time he was constantly engaged in scientific researches, many of which, such as his observations of gun-shot wounds, he put to excellent use in later life. On returning to England much improved in health in 1763, he entered at once upon his career as a London surgeon, and from that time forward his progress was a practically uninterrupted series of successes in his profession.

Hunter’s work on the study of the lymphatics was of great service to the medical profession. This important net-work of minute vessels distributed throughout the body had recently been made the object of much study, and various students, including Haller, had made extensive investigations since their discovery by Asellius. But Hunter, in 1758, was the first to discover the lymphatics in the neck of birds, although it was his brother William who advanced the theory that the function of these vessels was that of absorbents. One of John Hunter’s pupils, William Hewson (1739-1774), first gave an account, in 1768, of the lymphatics in reptiles and fishes, and added to his teacher’s investigations of the lymphatics in birds. These studies of the lymphatics have been regarded, perhaps with justice, as Hunter’s most valuable contributions to practical medicine.

In 1767 he met with an accident by which he suffered a rupture of the tendo Achillis–the large tendon that forms the attachment of the muscles of the calf to the heel. From observations of this accident, and subsequent experiments upon dogs, he laid the foundation for the now simple and effective operation for the cure of club feet and other deformities involving the tendons. In 1772 he moved into his residence at Earlscourt, Brompton, where he gathered about him a great menagerie of animals, birds, reptiles, insects, and fishes, which he used in his physiological and surgical experiments. Here he performed a countless number of experiments–more, probably, than “any man engaged in professional practice has ever conducted.” These experiments varied in nature from observations of the habits of bees and wasps to major surgical operations performed upon hedgehogs, dogs, leopards, etc. It is said that for fifteen years he kept a flock of geese for the sole purpose of studying the process of development in eggs.

Hunter began his first course of lectures in 1772, being forced to do this because he had been so repeatedly misquoted, and because he felt that he could better gauge his own knowledge in this way. Lecturing was a sore trial to him, as he was extremely diffident, and without writing out his lectures in advance he was scarcely able to speak at all. In this he presented a marked contrast to his brother William, who was a fluent and brilliant speaker. Hunter’s lectures were at best simple readings of the facts as he had written them, the diffident teacher seldom raising his eyes from his manuscript and rarely stopping until his complete lecture had been read through. His lectures were, therefore, instructive rather than interesting, as he used infinite care in preparing them; but appearing before his classes was so dreaded by him that he is said to have been in the habit of taking a half-drachm of laudanum before each lecture to nerve him for the ordeal. One is led to wonder by what name he shall designate that quality of mind that renders a bold and fearless surgeon like Hunter, who is undaunted in the face of hazardous and dangerous operations, a stumbling, halting, and “frightened” speaker before a little band of, at most, thirty young medical students. And yet this same thing is not unfrequently seen among the boldest surgeons.

Hunter’s Operation for the Cure of Aneurisms

It should be an object-lesson to those who, ignorantly or otherwise, preach against the painless vivisection as practised to-day, that by the sacrifice of a single deer in the cause of science Hunter discovered a fact in physiology that has been the means of saving thousands of human lives and thousands of human bodies from needless mutilation. We refer to the discovery of the “collateral circulation” of the blood, which led, among other things, to Hunter’s successful operation upon aneurisms.

Simply stated, every organ or muscle of the body is supplied by one large artery, whose main trunk distributes the blood into its lesser branches, and thence through the capillaries. Cutting off this main artery, it would seem, should cut off entirely the blood-supply to the particular organ which is supplied by this vessel; and until the time of Hunter’s demonstration this belief was held by most physiologists. But nature has made a provision for this possible stoppage of blood-supply from a single source, and has so arranged that some of the small arterial branches coming from the main supply-trunk are connected with other arterial branches coming from some other supply-trunk. Under normal conditions the main arterial trunks supply their respective organs, the little connecting arterioles playing an insignificant part. But let the main supply-trunk be cut off or stopped for whatever reason, and a remarkable thing takes place. The little connecting branches begin at once to enlarge and draw blood from the neighboring uninjured supply-trunk, This enlargement continues until at last a new route for the circulation has been established, the organ no longer depending on the now defunct original arterial trunk, but getting on as well as before by this “collateral” circulation that has been established.

The thorough understanding of this collateral circulation is one of the most important steps in surgery, for until it was discovered amputations were thought necessary in such cases as those involving the artery supplying a leg or arm, since it was supposed that, the artery being stopped, death of the limb and the subsequent necessity for amputation were sure to follow. Hunter solved this problem by a single operation upon a deer, and his practicality as a surgeon led him soon after to apply this knowledge to a certain class of surgical cases in a most revolutionary and satisfactory manner.

What led to Hunter’s far-reaching discovery was his investigation as to the cause of the growth of the antlers of the deer. Wishing to ascertain just what part the blood-supply on the opposite sides of the neck played in the process of development, or, perhaps more correctly, to see what effect cutting off the main blood-supply would have, Hunter had one of the deer of Richmond Park caught and tied, while he placed a ligature around one of the carotid arteries–one of the two principal arteries that supply the head with blood. He observed that shortly after this the antler (which was only half grown and consequently very vascular) on the side of the obliterated artery became cold to the touch–from the lack of warmth-giving blood. There was nothing unexpected in this, and Hunter thought nothing of it until a few days later, when he found, to his surprise, that the antler had become as warm as its fellow, and was apparently increasing in size. Puzzled as to how this could be, and suspecting that in some way his ligature around the artery had not been effective, he ordered the deer killed, and on examination was astonished to find that while his ligature had completely shut off the blood-supply from the source of that carotid artery, the smaller arteries had become enlarged so as to supply the antler with blood as well as ever, only by a different route.

Hunter soon had a chance to make a practical application of the knowledge thus acquired. This was a case of popliteal aneurism, operations for which had heretofore proved pretty uniformly fatal. An aneurism, as is generally understood, is an enlargement of a certain part of an artery, this enlargement sometimes becoming of enormous size, full of palpitating blood, and likely to rupture with fatal results at any time. If by any means the blood can be allowed to remain quiet for even a few hours in this aneurism it will form a clot, contract, and finally be absorbed and disappear without any evil results. The problem of keeping the blood quiet, with the heart continually driving it through the vessel, is not a simple one, and in Hunter’s time was considered so insurmountable that some surgeons advocated amputation of any member having an aneurism, while others cut down upon the tumor itself and attempted to tie off the artery above and below. The first of these operations maimed the patient for life, while the second was likely to prove fatal.

In pondering over what he had learned about collateral circulation and the time required for it to become fully established, Hunter conceived the idea that if the blood-supply was cut off from above the aneurism, thus temporarily preventing the ceaseless pulsations from the heart, this blood would coagulate and form a clot before the collateral circulation could become established or could affect it. The patient upon whom he performed his now celebrated operation was afflicted with a popliteal aneurism–that is, the aneurism was located on the large popliteal artery just behind the knee-joint. Hunter, therefore, tied off the femoral, or main supplying artery in the thigh, a little distance above the aneurism. The operation was entirely successful, and in six weeks’ time the patient was able to leave the hospital, and with two sound limbs. Naturally the simplicity and success of this operation aroused the attention of Europe, and, alone, would have made the name of Hunter immortal in the annals of surgery. The operation has ever since been called the “Hunterian” operation for aneurism, but there is reason to believe that Dominique Anel (born about 1679) performed a somewhat similar operation several years earlier. It is probable, however, that Hunter had never heard of this work of Anel, and that his operation was the outcome of his own independent reasoning from the facts he had learned about collateral circulation. Furthermore, Hunter’s mode of operation was a much better one than Anel’s, and, while Anel’s must claim priority, the credit of making it widely known will always be Hunter’s.

The great services of Hunter were recognized both at home and abroad, and honors and positions of honor and responsibility were given him. In 1776 he was appointed surgeon-extraordinary to the king; in 1783 he was elected a member of the Royal Society of Medicine and of the Royal Academy of Surgery at Paris; in 1786 he became deputy surgeon-general of the army; and in 1790 he was appointed surgeon-general and inspector-general of hospitals. All these positions he filled with credit, and he was actively engaged in his tireless pursuit of knowledge and in discharging his many duties when in October, 1793, he was stricken while addressing some colleagues, and fell dead in the arms of a fellow-physician.


Hunter’s great rival among contemporary physiologists was the Italian Lazzaro Spallanzani (1729-1799), one of the most picturesque figures in the history of science. He was not educated either as a scientist or physician, devoting, himself at first to philosophy and the languages, afterwards studying law, and later taking orders. But he was a keen observer of nature and of a questioning and investigating mind, so that he is remembered now chiefly for his discoveries and investigations in the biological sciences. One important demonstration was his controversion of the theory of abiogenesis, or “spontaneous generation,” as propounded by Needham and Buffon. At the time of Needham’s experiments it had long been observed that when animal or vegetable matter had lain in water for a little time–long enough for it to begin to undergo decomposition–the water became filled with microscopic creatures, the “infusoria animalculis.” This would tend to show, either that the water or the animal or vegetable substance contained the “germs” of these minute organisms, or else that they were generated spontaneously. It was known that boiling killed these animalcules, and Needham agreed, therefore, that if he first heated the meat or vegetables, and also the water containing them, and then placed them in hermetically scaled jars–if he did this, and still the animalcules made their appearance, it would be proof-positive that they had been generated spontaneously. Accordingly be made numerous experiments, always with the same results–that after a few days the water was found to swarm with the microscopic creatures. The thing seemed proven beyond question–providing, of course, that there had been no slips in the experiments.

But Abbe Spallanzani thought that he detected such slips in Needham’s experiment. The possibility of such slips might come in several ways: the contents of the jar might not have been boiled for a sufficient length of time to kill all the germs, or the air might not have been excluded completely by the sealing process. To cover both these contingencies, Spallanzani first hermetically sealed the glass vessels and then boiled them for three-quarters of an hour. Under these circumstances no animalcules ever made their appearance–a conclusive demonstration that rendered Needham’s grounds for his theory at once untenable.[2]

Allied to these studies of spontaneous generation were Spallanzani’s experiments and observations on the physiological processes of generation among higher animals. He experimented with frogs, tortoises, and dogs; and settled beyond question the function of the ovum and spermatozoon. Unfortunately he misinterpreted the part played by the spermatozoa in believing that their surrounding fluid was equally active in the fertilizing process, and it was not until some forty years later (1824) that Dumas corrected this error.


Among the most interesting researches of Spallanzani were his experiments to prove that digestion, as carried on in the stomach, is a chemical process. In this he demonstrated, as Rene Reaumur had attempted to demonstrate, that digestion could be carried on outside the walls of the stomach as an ordinary chemical reaction, using the gastric juice as the reagent for performing the experiment. The question as to whether the stomach acted as a grinding or triturating organ, rather than as a receptacle for chemical action, had been settled by Reaumur and was no longer a question of general dispute. Reaumur had demonstrated conclusively that digestion would take place in the stomach in the same manner and the same time if the substance to be digested was protected from the peristalic movements of the stomach and subjected to the action of the gastric juice only. He did this by introducing the substances to be digested into the stomach in tubes, and thus protected so that while the juices of the stomach could act upon them freely they would not be affected by any movements of the organ.

Following up these experiments, he attempted to show that digestion could take place outside the body as well as in it, as it certainly should if it were a purely chemical process. He collected quantities of gastric juice, and placing it in suitable vessels containing crushed grain or flesh, kept the mixture at about the temperature of the body for several hours. After repeated experiments of this kind, apparently conducted with great care, Reaumur reached the conclusion that “the gastric juice has no more effect out of the living body in dissolving or digesting the food than water, mucilage, milk, or any other bland fluid.”[3] Just why all of these experiments failed to demonstrate a fact so simple does not appear; but to Spallanzani, at least, they were by no means conclusive, and he proceeded to elaborate upon the experiments of Reaumur. He made his experiments in scaled tubes exposed to a certain degree of heat, and showed conclusively that the chemical process does go on, even when the food and gastric juice are removed from their natural environment in the stomach. In this he was opposed by many physiologists, among them John Hunter, but the truth of his demonstrations could not be shaken, and in later years we find Hunter himself completing Spallanzani’s experiments by his studies of the post-mortem action of the gastric juice upon the stomach walls.

That Spallanzani’s and Hunter’s theories of the action of the gastric juice were not at once universally accepted is shown by an essay written by a learned physician in 1834. In speaking of some of Spallanzani’s demonstrations, he writes: “In some of the experiments, in order to give the flesh or grains steeped in the gastric juice the same temperature with the body, the phials were introduced under the armpits. But this is not a fair mode of ascertaining the effects of the gastric juice out of the body; for the influence which life may be supposed to have on the solution of the food would be secured in this case. The affinities connected with life would extend to substances in contact with any part of the system: substances placed under the armpits are not placed at least in the same circumstances with those unconnected with a living animal.” But just how this writer reaches the conclusion that “the experiments of Reaumur and Spallanzani give no evidence that the gastric juice has any peculiar influence more than water or any other bland fluid in digesting the food”[4] is difficult to understand.

The concluding touches were given to the new theory of digestion by John Hunter, who, as we have seen, at first opposed Spallanzani, but who finally became an ardent champion of the chemical theory. Hunter now carried Spallanzani’s experiments further and proved the action of the digestive fluids after death. For many years anatomists had been puzzled by pathological lesion of the stomach, found post mortem, when no symptoms of any disorder of the stomach had been evinced during life. Hunter rightly conceived that these lesions were caused by the action of the gastric juice, which, while unable to act upon the living tissue, continued its action chemically after death, thus digesting the walls of the stomach in which it had been formed. And, as usual with his observations, be turned this discovery to practical use in accounting for certain phenomena of digestion. The following account of the stomach being digested after death was written by Hunter at the desire of Sir John Pringle, when he was president of the Royal Society, and the circumstance which led to this is as follows: “I was opening, in his presence, the body of a patient of his own, where the stomach was in part dissolved, which appeared to him very unaccountable, as there had been no previous symptom that could have led him to suspect any disease in the stomach. I took that opportunity of giving him my ideas respecting it, and told him that I had long been making experiments on digestion, and considered this as one of the facts which proved a converting power in the gastric juice. . . . There are a great many powers in nature which the living principle does not enable the animal matter, with which it is combined, to resist–viz., the mechanical and most of the strongest chemical solvents. It renders it, however, capable of resisting the powers of fermentation, digestion, and perhaps several others, which are well known to act on the same matter when deprived of the living principle and entirely to decompose it. “

Hunter concludes his paper with the following paragraph: “These appearances throw considerable light on the principle of digestion, and show that it is neither a mechanical power, nor contractions of the stomach, nor heat, but something secreted in the coats of the stomach, and thrown into its cavity, which there animalizes the food or assimilates it to the nature of the blood. The power of this juice is confined or limited to certain substances, especially of the vegetable and animal kingdoms; and although this menstruum is capable of acting independently of the stomach, yet it is indebted to that viscus for its continuance.[5]


It is a curious commentary on the crude notions of mechanics of previous generations that it should have been necessary to prove by experiment that the thin, almost membranous stomach of a mammal has not the power to pulverize, by mere attrition, the foods that are taken into it. However, the proof was now for the first time forthcoming, and the question of the general character of the function of digestion was forever set at rest. Almost simultaneously with this great advance, corresponding progress was made in an allied field: the mysteries of respiration were at last cleared up, thanks to the new knowledge of chemistry. The solution of the problem followed almost as a matter of course upon the advances of that science in the latter part of the century. Hitherto no one since Mayow, of the previous century, whose flash of insight had been strangely overlooked and forgotten, had even vaguely surmised the true function of the lungs. The great Boerhaave had supposed that respiration is chiefly important as an aid to the circulation of the blood; his great pupil, Haller, had believed to the day of his death in 1777 that the main purpose of the function is to form the voice. No genius could hope to fathom the mystery of the lungs so long as air was supposed to be a simple element, serving a mere mechanical purpose in the economy of the earth.

But the discovery of oxygen gave the clew, and very soon all the chemists were testing the air that came from the lungs–Dr. Priestley, as usual, being in the van. His initial experiments were made in 1777, and from the outset the problem was as good as solved. Other experimenters confirmed his results in all their essentials–notably Scheele and Lavoisier and Spallanzani and Davy. It was clearly established that there is chemical action in the contact of the air with the tissue of the lungs; that some of the oxygen of the air disappears, and that carbonic-acid gas is added to the inspired air. It was shown, too, that the blood, having come in contact with the air, is changed from black to red in color. These essentials were not in dispute from the first. But as to just what chemical changes caused these results was the subject of controversy. Whether, for example, oxygen is actually absorbed into the blood, or whether it merely unites with carbon given off from the blood, was long in dispute.

Each of the main disputants was biased by his own particular views as to the moot points of chemistry. Lavoisier, for example, believed oxygen gas to be composed of a metal oxygen combined with the alleged element heat; Dr. Priestley thought it a compound of positive electricity and phlogiston; and Humphry Davy, when he entered the lists a little later, supposed it to be a compound of oxygen and light. Such mistaken notions naturally complicated matters and delayed a complete understanding of the chemical processes of respiration. It was some time, too, before the idea gained acceptance that the most important chemical changes do not occur in the lungs themselves, but in the ultimate tissues. Indeed, the matter was not clearly settled at the close of the century. Nevertheless, the problem of respiration had been solved in its essentials. Moreover, the vastly important fact had been established that a process essentially identical with respiration is necessary to the existence not only of all creatures supplied with lungs, but to fishes, insects, and even vegetables–in short, to every kind of living organism.


Some interesting experiments regarding vegetable respiration were made just at the close of the century by Erasmus Darwin, and recorded in his Botanic Garden as a foot-note to the verse:

“While spread in air the leaves respiring play.”

These notes are worth quoting at some length, as they give a clear idea of the physiological doctrines of the time (1799), while taking advance ground as to the specific matter in question:

“There have been various opinions,” Darwin says, “concerning the use of the leaves of plants in the vegetable economy. Some have contended that they are perspiratory organs. This does not seem probable from an experiment of Dr. Hales, Vegetable Statics, p. 30. He, found, by cutting off branches of trees with apples on them and taking off the leaves, that an apple exhaled about as much as two leaves the surfaces of which were nearly equal to the apple; whence it would appear that apples have as good a claim to be termed perspiratory organs as leaves. Others have believed them excretory organs of excrementitious juices, but as the vapor exhaled from vegetables has no taste, this idea is no more probable than the other; add to this that in most weathers they do not appear to perspire or exhale at all.

“The internal surface of the lungs or air-vessels in men is said to be equal to the external surface of the whole body, or almost fifteen square feet; on this surface the blood is exposed to the influence of the respired air through the medium, however, of a thin pellicle; by this exposure to the air it has its color changed from deep red to bright scarlet, and acquires something so necessary to the existence of life that we can live scarcely a minute without this wonderful process.

“The analogy between the leaves of plants and the lungs or gills of animals seems to embrace so many circumstances that we can scarcely withhold our consent to their performing similar offices.

“1. The great surface of leaves compared to that of the trunk and branches of trees is such that it would seem to be an organ well adapted for the purpose of exposing the vegetable juices to the influence of the air; this, however, we shall see afterwards is probably performed only by their upper surfaces, yet even in this case the surface of the leaves in general bear a greater proportion to the surface of the tree than the lungs of animals to their external surfaces.

“2. In the lung of animals the blood, after having been exposed to the air in the extremities of the pulmonary artery, is changed in color from deep red to bright scarlet, and certainly in some of its essential properties it is then collected by the pulmonary vein and returned to the heart. To show a similarity of circumstances in the leaves of plants, the following experiment was made, June 24, 1781. A stalk with leaves and seed-vessels of large spurge (Euphorbia helioscopia) had been several days placed in a decoction of madder (Rubia tinctorum) so that the lower part of the stem and two of the undermost leaves were immersed in it. After having washed the immersed leaves in clear water I could readily discover the color of the madder passing along the middle rib of each leaf. The red artery was beautifully visible on the under and on the upper surface of the leaf; but on the upper side many red branches were seen going from it to the extremities of the leaf, which on the other side were not visible except by looking through it against the light. On this under side a system of branching vessels carrying a pale milky fluid were seen coming from the extremities of the leaf, and covering the whole under side of it, and joining two large veins, one on each side of the red artery in the middle rib of the leaf, and along with it descending to the foot-stalk or petiole. On slitting one of these leaves with scissors, and having a magnifying-glass ready, the milky blood was seen oozing out of the returning veins on each side of the red artery in the middle rib, but none of the red fluid from the artery.

“All these appearances were more easily seen in a leaf of Picris treated in the same manner; for in this milky plant the stems and middle rib of the leaves are sometimes naturally colored reddish, and hence the color of the madder seemed to pass farther into the ramifications of their leaf-arteries, and was there beautifully visible with the returning branches of milky veins on each side.”

Darwin now goes on to draw an incorrect inference from his observations:

“3. From these experiments,” he says, “the upper surface of the leaf appeared to be the immediate organ of respiration, because the colored fluid was carried to the extremities of the leaf by vessels most conspicuous on the upper surface, and there changed into a milky fluid, which is the blood of the plant, and then returned by concomitant veins on the under surface, which were seen to ooze when divided with scissors, and which, in Picris, particularly, render the under surface of the leaves greatly whiter than the upper one.”

But in point of fact, as studies of a later generation were to show, it is the under surface of the leaf that is most abundantly provided with stomata, or “breathing-pores.” From the stand-point of this later knowledge, it is of interest to follow our author a little farther, to illustrate yet more fully the possibility of combining correct observations with a faulty inference.

“4. As the upper surface of leaves constitutes the organ of respiration, on which the sap is exposed in the termination of arteries beneath a thin pellicle to the action of the atmosphere, these surfaces in many plants strongly repel moisture, as cabbage leaves, whence the particles of rain lying over their surfaces without touching them, as observed by Mr. Melville (Essays Literary and Philosophical: Edinburgh), have the appearance of globules of quicksilver. And hence leaves with the upper surfaces on water wither as soon as in the dry air, but continue green for many days if placed with the under surface on water, as appears in the experiments of Monsieur Bonnet (Usage des Feuilles). Hence some aquatic plants, as the water-lily (Nymphoea), have the lower sides floating on the water, while the upper surfaces remain dry in the air.

“5. As those insects which have many spiracula, or breathing apertures, as wasps and flies, are immediately suffocated by pouring oil upon them, I carefully covered with oil the surfaces of several leaves of phlomis, of Portugal laurel, and balsams, and though it would not regularly adhere, I found them all die in a day or two.

“It must be added that many leaves are furnished with muscles about their foot-stalks, to turn their surfaces to the air or light, as mimosa or Hedysarum gyrans. From all these analogies I think there can be no doubt but that leaves of trees are their lungs, giving out a phlogistic material to the atmosphere, and absorbing oxygen, or vital air.

“6. The great use of light to vegetation would appear from this theory to be by disengaging vital air from the water which they perspire, and thence to facilitate its union with their blood exposed beneath the thin surface of their leaves; since when pure air is thus applied it is probable that it can be more readily absorbed. Hence, in the curious experiments of Dr. Priestley and Mr. Ingenhouz, some plants purified less air than others–that is, they perspired less in the sunshine; and Mr. Scheele found that by putting peas into water which about half covered them they converted the vital air into fixed air, or carbonic-acid gas, in the same manner as in animal respiration.

“7. The circulation in the lungs or leaves of plants is very similar to that of fish. In fish the blood, after having passed through their gills, does not return to the heart as from the lungs of air-breathing animals, but the pulmonary vein taking the structure of an artery after having received the blood from the gills, which there gains a more florid color, distributes it to the other parts of their bodies. The same structure occurs in the livers of fish, whence we see in those animals two circulations independent of the power of the heart–viz., that beginning at the termination of the veins of the gills and branching through the muscles, and that which passes through the liver; both which are carried on by the action of those respective arteries and veins.”[6]

Darwin is here a trifle fanciful in forcing the analogy between plants and animals. The circulatory system of plants is really not quite so elaborately comparable to that of fishes as he supposed. But the all-important idea of the uniformity underlying the seeming diversity of Nature is here exemplified, as elsewhere in the writings of Erasmus Darwin; and, more specifically, a clear grasp of the essentials of the function of respiration is fully demonstrated.


Several causes conspired to make exploration all the fashion during the closing epoch of the eighteenth century. New aid to the navigator had been furnished by the perfected compass and quadrant, and by the invention of the chronometer; medical science had banished scurvy, which hitherto had been a perpetual menace to the voyager; and, above all, the restless spirit of the age impelled the venturesome to seek novelty in fields altogether new. Some started for the pole, others tried for a northeast or northwest passage to India, yet others sought the great fictitious antarctic continent told of by tradition. All these of course failed of their immediate purpose, but they added much to the world’s store of knowledge and its fund of travellers’ tales.

Among all these tales none was more remarkable than those which told of strange living creatures found in antipodal lands. And here, as did not happen in every field, the narratives were often substantiated by the exhibition of specimens that admitted no question. Many a company of explorers returned more or less laden with such trophies from the animal and vegetable kingdoms, to the mingled astonishment, delight, and bewilderment of the closet naturalists. The followers of Linnaeus in the “golden age of natural history,” a few decades before, had increased the number of known species of fishes to about four hundred, of birds to one thousand, of insects to three thousand, and of plants to ten thousand. But now these sudden accessions from new territories doubled the figure for plants, tripled it for fish and birds, and brought the number of described insects above twenty thousand. Naturally enough, this wealth of new material was sorely puzzling to the classifiers. The more discerning began to see that the artificial system of Linnaeus, wonderful and useful as it had been, must be advanced upon before the new material could be satisfactorily disposed of. The way to a more natural system, based on less arbitrary signs, had been pointed out by Jussieu in botany, but the zoologists were not prepared to make headway towards such a system until they should gain a wider understanding of the organisms with which they had to deal through comprehensive studies of anatomy. Such studies of individual forms in their relations to the entire scale of organic beings were pursued in these last decades of the century, but though two or three most important generalizations were achieved (notably Kaspar Wolff’s conception of the cell as the basis of organic life, and Goethe’s all-important doctrine of metamorphosis of parts), yet, as a whole, the work of the anatomists of the period was germinative rather than fruit-bearing. Bichat’s volumes, telling of the recognition of the fundamental tissues of the body, did not begin to appear till the last year of the century. The announcement by Cuvier of the doctrine of correlation of parts bears the same date, but in general the studies of this great naturalist, which in due time were to stamp him as the successor of Linnaeus, were as yet only fairly begun.



We have seen that the focal points of the physiological world towards the close of the eighteenth century were Italy and England, but when Spallanzani and Hunter passed away the scene shifted to France. The time was peculiarly propitious, as the recent advances in many lines of science had brought fresh data for the student of animal life which were in need of classification, and, as several minds capable of such a task were in the field, it was natural that great generalizations should have come to be quite the fashion. Thus it was that Cuvier came forward with a brand-new classification of the animal kingdom, establishing four great types of being, which he called vertebrates, mollusks, articulates, and radiates. Lamarck had shortly before established the broad distinction between animals with and those without a backbone; Cuvier’s Classification divided the latter–the invertebrates–into three minor groups. And this division, familiar ever since to all students of zoology, has only in very recent years been supplanted, and then not by revolution, but by a further division, which the elaborate recent studies of lower forms of life seemed to make desirable.

In the course of those studies of comparative anatomy which led to his new classification, Cuvier’s attention was called constantly to the peculiar co-ordination of parts in each individual organism. Thus an animal with sharp talons for catching living prey–as a member of the cat tribe–has also sharp teeth, adapted for tearing up the flesh of its victim, and a particular type of stomach, quite different from that of herbivorous creatures. This adaptation of all the parts of the animal to one another extends to the most diverse parts of the organism, and enables the skilled anatomist, from the observation of a single typical part, to draw inferences as to the structure of the entire animal–a fact which was of vast aid to Cuvier in his studies of paleontology. It did not enable Cuvier, nor does it enable any one else, to reconstruct fully the extinct animal from observation of a single bone, as has sometimes been asserted, but what it really does establish, in the hands of an expert, is sufficiently astonishing.

“While the study of the fossil remains of the greater quadrupeds is more satisfactory,” he writes, “by the clear results which it affords, than that of the remains of other animals found in a fossil state, it is also complicated with greater and more numerous difficulties. Fossil shells are usually found quite entire, and retaining all the characters requisite for comparing them with the specimens contained in collections of natural history, or represented in the works of naturalists. Even the skeletons of fishes are found more or less entire, so that the general forms of their bodies can, for the most part, be ascertained, and usually, at least, their generic and specific characters are determinable, as these are mostly drawn from their solid parts. In quadrupeds, on the contrary, even when their entire skeletons are found, there is great difficulty in discovering their distinguishing characters, as these are chiefly founded upon their hairs and colors and other marks which have disappeared previous to their incrustation. It is also very rare to find any fossil skeletons of quadrupeds in any degree approaching to a complete state, as the strata for the most part only contain separate bones, scattered confusedly and almost always broken and reduced to fragments, which are the only means left to naturalists for ascertaining the species or genera to which they have belonged.

“Fortunately comparative anatomy, when thoroughly understood, enables us to surmount all these difficulties, as a careful application of its principles instructs us in the correspondences and dissimilarities of the forms of organized bodies of different kinds, by which each may be rigorously ascertained from almost every fragment of its various parts and organs.

“Every organized individual forms an entire system of its own, all the parts of which naturally correspond, and concur to produce a certain definite purpose, by reciprocal reaction, or by combining towards the same end. Hence none of these separate parts can change their forms without a corresponding change in the other parts of the same animal, and consequently each of these parts, taken separately, indicates all the other parts to which it has belonged. Thus, as I have elsewhere shown, if the viscera of an animal are so organized as only to be fitted for the digestion of recent flesh, it is also requisite that the jaws should be so constructed as to fit them for devouring prey; the claws must be constructed for seizing and tearing it to pieces; the teeth for cutting and dividing its flesh; the entire system of the limbs, or organs of motion, for pursuing and overtaking it; and the organs of sense for discovering it at a distance. Nature must also have endowed the brain of the animal with instincts sufficient for concealing itself and for laying plans to catch its necessary victims. . . . . . . . . .

“To enable the animal to carry off its prey when seized, a corresponding force is requisite in the muscles which elevate the head, and this necessarily gives rise to a determinate form of the vertebrae to which these muscles are attached and of the occiput into which they are inserted. In order that the teeth of a carnivorous animal may be able to cut the flesh, they require to be sharp, more or less so in proportion to the greater or less quantity of flesh that they have to cut. It is requisite that their roots should be solid and strong, in proportion to the quantity and size of the bones which they have to break to pieces. The whole of these circumstances must necessarily influence the development and form of all the parts which contribute to move the jaws. . . . . . . . . .

After these observations, it will be easily seen that similar conclusions may be drawn with respect to the limbs of carnivorous animals, which require particular conformations to fit them for rapidity of motion in general; and that similar considerations must influence the forms and connections of the vertebrae and other bones constituting the trunk of the body, to fit them for flexibility and readiness of motion in all directions. The bones also of the nose, of the orbit, and of the ears require certain forms and structures to fit them for giving perfection to the senses of smell, sight, and hearing, so necessary to animals of prey. In short, the shape and structure of the teeth regulate the forms of the condyle, of the shoulder-blade, and of the claws, in the same manner as the equation of a curve regulates all its other properties; and as in regard to any particular curve all its properties may be ascertained by assuming each separate property as the foundation of a particular equation, in the same manner a claw, a shoulder-blade, a condyle, a leg or arm bone, or any other bone separately considered, enables us to discover the description of teeth to which they have belonged; and so also reciprocally we may determine the forms of the other bones from the teeth. Thus commencing our investigations by a careful survey of any one bone by itself, a person who is sufficiently master of the laws of organic structure may, as it were, reconstruct the whole animal to which that bone belonged.”[1]

We have already pointed out that no one is quite able to perform the necromantic feat suggested in the last sentence; but the exaggeration is pardonable in the enthusiast to whom the principle meant so much and in whose hands it extended so far.

Of course this entire principle, in its broad outlines, is something with which every student of anatomy had been familiar from the time when anatomy was first studied, but the full expression of the “law of co-ordination,” as Cuvier called it, had never been explicitly made before; and, notwithstanding its seeming obviousness, the exposition which Cuvier made of it in the introduction to his classical work on comparative anatomy, which was published during the first decade of the nineteenth century, ranks as a great discovery. It is one of those generalizations which serve as guideposts to other discoveries.


Much the same thing may be said of another generalization regarding the animal body, which the brilliant young French physician Marie Francois Bichat made in calling attention to the fact that each vertebrate organism, including man, has really two quite different sets of organs–one set under volitional control, and serving the end of locomotion, the other removed from volitional control, and serving the ends of the “vital processes” of digestion, assimilation, and the like. He called these sets of organs the animal system and the organic system, respectively. The division thus pointed out was not quite new, for Grimaud, professor of physiology in the University of Montpellier, had earlier made what was substantially the same classification of the functions into “internal or digestive and external or locomotive”; but it was Bichat’s exposition that gave currency to the idea.

Far more important, however, was another classification which Bichat put forward in his work on anatomy, published just at the beginning of the last century. This was the division of all animal structures into what Bichat called tissues, and the pointing out that there are really only a few kinds of these in the body, making up all the diverse organs. Thus muscular organs form one system; membranous organs another; glandular organs a third; the vascular mechanism a fourth, and so on. The distinction is so obvious that it seems rather difficult to conceive that it could have been overlooked by the earliest anatomists; but, in point of fact, it is only obvious because now it has been familiarly taught for almost a century. It had never been given explicit expression before the time of Bichat, though it is said that Bichat himself was somewhat indebted for it to his master, Desault, and to the famous alienist Pinel.

However that may be, it is certain that all subsequent anatomists have found Bichat’s classification of the tissues of the utmost value in their studies of the animal functions. Subsequent advances were to show that the distinction between the various tissues is not really so fundamental as Bichat supposed, but that takes nothing from the practical value of the famous classification.

It was but a step from this scientific classification of tissues to a similar classification of the diseases affecting them, and this was one of the greatest steps towards placing medicine on the plane of an exact science. This subject of these branches completely fascinated Bichat, and he exclaimed, enthusiastically: “Take away some fevers and nervous trouble, and all else belongs to the kingdom of pathological anatomy.” But out of this enthusiasm came great results. Bichat practised as he preached, and, believing that it was only possible to understand disease by observing the symptoms carefully at the bedside, and, if the disease terminated fatally, by post-mortem examination, he was so arduous in his pursuit of knowledge that within a period of less than six months he had made over six hundred autopsies–a record that has seldom, if ever, been equalled. Nor were his efforts fruitless, as a single example will suffice to show. By his examinations he was able to prove that diseases of the chest, which had formerly been classed under the indefinite name “peripneumonia,” might involve three different structures, the pleural sac covering the lungs, the lung itself, and the bronchial tubes, the diseases affecting these organs being known respectively as pleuritis, pneumonia, and bronchitis, each one differing from the others as to prognosis and treatment. The advantage of such an exact classification needs no demonstration.


At the same time when these broad macroscopical distinctions were being drawn there were other workers who were striving to go even deeper into the intricacies of the animal mechanism with the aid of the microscope. This undertaking, however, was beset with very great optical difficulties, and for a long time little advance was made upon the work of preceding generations. Two great optical barriers, known technically as spherical and chromatic aberration–the one due to a failure of the rays of light to fall all in one plane when focalized through a lens, the other due to the dispersive action of the lens in breaking the white light into prismatic colors–confronted the makers of microscopic lenses, and seemed all but insuperable. The making of achromatic lenses for telescopes had been accomplished, it is true, by Dolland in the previous century, by the union of lenses of crown glass with those of flint glass, these two materials having different indices of refraction and dispersion. But, aside from the mechanical difficulties which arise when the lens is of the minute dimensions required for use with the microscope, other perplexities are introduced by the fact that the use of a wide pencil of light is a desideratum, in order to gain sufficient illumination when large magnification is to be secured.

In the attempt to overcome those difficulties, the foremost physical philosophers of the time came to the aid of the best opticians. Very early in the century, Dr. (afterwards Sir David) Brewster, the renowned Scotch physicist, suggested that certain advantages might accrue from the use of such gems as have high refractive and low dispersive indices, in place of lenses made of glass. Accordingly lenses were made of diamond, of sapphire, and so on, and with some measure of success. But in 1812 a much more important innovation was introduced by Dr. William Hyde Wollaston, one of the greatest and most versatile, and, since the death of Cavendish, by far the most eccentric of English natural philosophers. This was the suggestion to use two plano-convex lenses, placed at a prescribed distance apart, in lieu of the single double-convex lens generally used. This combination largely overcame the spherical aberration, and it gained immediate fame as the “Wollaston doublet.”

To obviate loss of light in such a doublet from increase of reflecting surfaces, Dr. Brewster suggested filling the interspace between the two lenses with a cement having the same index of refraction as the lenses themselves–an improvement of manifest advantage. An improvement yet more important was made by Dr. Wollaston himself in the introduction of the diaphragm to limit the field of vision between the lenses, instead of in front of the anterior lens. A pair of lenses thus equipped Dr. Wollaston called the periscopic microscope. Dr. Brewster suggested that in such a lens the same object might be attained with greater ease by grinding an equatorial groove about a thick or globular lens and filling the groove with an opaque cement. This arrangement found much favor, and came subsequently to be known as a Coddington lens, though Mr. Coddington laid no claim to being its inventor.

Sir John Herschel, another of the very great physicists of the time, also gave attention to the problem of improving the microscope, and in 1821 he introduced what was called an aplanatic combination of lenses, in which, as the name implies, the spherical aberration was largely done away with. It was thought that the use of this Herschel aplanatic combination as an eyepiece, combined with the Wollaston doublet for the objective, came as near perfection as the compound microscope was likely soon to come. But in reality the instrument thus constructed, though doubtless superior to any predecessor, was so defective that for practical purposes the simple microscope, such as the doublet or the Coddington, was preferable to the more complicated one.

Many opticians, indeed, quite despaired of ever being able to make a satisfactory refracting compound microscope, and some of them had taken up anew Sir Isaac Newton’s suggestion in reference to a reflecting microscope. In particular, Professor Giovanni Battista Amici, a very famous mathematician and practical optician of Modena, succeeded in constructing a reflecting microscope which was said to be superior to any compound microscope of the time, though the events of the ensuing years were destined to rob it of all but historical value. For there were others, fortunately, who did not despair of the possibilities of the refracting microscope, and their efforts were destined before long to be crowned with a degree of success not even dreamed of by any preceding generation.

The man to whom chief credit is due for directing those final steps that made the compound microscope a practical implement instead of a scientific toy was the English amateur optician Joseph Jackson Lister. Combining mathematical knowledge with mechanical ingenuity, and having the practical aid of the celebrated optician Tulley, he devised formulae for the combination of lenses of crown glass with others of flint glass, so adjusted that the refractive errors of one were corrected or compensated by the other, with the result of producing lenses of hitherto unequalled powers of definition; lenses capable of showing an image highly magnified, yet relatively free from those distortions and fringes of color that had heretofore been so disastrous to true interpretation of magnified structures.

Lister had begun his studies of the lens in 1824, but it was not until 1830 that he contributed to the Royal Society the famous paper detailing his theories and experiments. Soon after this various continental opticians who had long been working along similar lines took the matter up, and their expositions, in particular that of Amici, introduced the improved compound microscope to the attention of microscopists everywhere. And it required but the most casual trial to convince the experienced observers that a new implement of scientific research had been placed in their hands which carried them a long step nearer the observation of the intimate physical processes which lie at the foundation of vital phenomena. For the physiologist this perfection of the compound microscope had the same significance that the, discovery of America had for the fifteenth-century geographers–it promised a veritable world of utterly novel revelations. Nor was the fulfilment of that promise long delayed.

Indeed, so numerous and so important were the discoveries now made in the realm of minute anatomy that the rise of histology to the rank of an independent science may be said to date from this period. Hitherto, ever since the discovery of magnifying-glasses, there had been here and there a man, such as Leuwenhoek or Malpighi, gifted with exceptional vision, and perhaps unusually happy in his conjectures, who made important contributions to the knowledge of the minute structure of organic tissues; but now of a sudden it became possible for the veriest tyro to confirm or refute the laborious observations of these pioneers, while the skilled observer could step easily beyond the barriers of vision that hitherto were quite impassable. And so, naturally enough, the physiologists of the fourth decade of the nineteenth century rushed as eagerly into the new realm of the microscope as, for example, their successors of to-day are exploring the realm of the X-ray.

Lister himself, who had become an eager interrogator of the instrument he had perfected, made many important discoveries, the most notable being his final settlement of the long-mooted question as to the true form of the red corpuscles of the human blood. In reality, as everybody knows nowadays, these are biconcave disks, but owing to their peculiar figure it is easily possible to misinterpret the appearances they present when seen through a poor lens, and though Dr. Thomas Young and various other observers had come very near the truth regarding them, unanimity of opinion was possible only after the verdict of the perfected microscope was given.

These blood corpuscles are so infinitesimal in size that something like five millions of them are found in each cubic millimetre of the blood, yet they are isolated particles, each having, so to speak, its own personality. This, of course, had been known to microscopists since the days of the earliest lenses. It had been noticed, too, by here and there an observer, that certain of the solid tissues seemed to present something of a granular texture, as if they, too, in their ultimate constitution, were made up of particles. And now, as better and better lenses were constructed, this idea gained ground constantly, though for a time no one saw its full significance. In the case of vegetable tissues, indeed, the fact that little particles encased a membranous covering, and called cells, are the ultimate visible units of structure had long been known. But it was supposed that animal tissues differed radically from this construction. The elementary particles of vegetables “were regarded to a certain extent as individuals which composed the entire plant, while, on the other hand, no such view was taken of the elementary parts of animals.”


In the year 1833 a further insight into the nature of the ultimate particles of plants was gained through the observation of the English microscopist Robert Brown, who, in the course of his microscopic studies of the epidermis of orchids, discovered in the cells “an opaque spot,” which he named the nucleus. Doubtless the same “spot” had been seen often enough before by other observers, but Brown was the first to recognize it as a component part of the vegetable cell and to give it a name.

“I shall conclude my observations on Orchideae,” said Brown, “with a notice of some points of their general structure, which chiefly relate to the cellular tissue. In each cell of the epidermis of a great part of this family, especially of those with membranous leaves, a single circular areola, generally somewhat more opaque than, the membrane of the cell, is observable. This areola, which is more or less distinctly granular, is slightly convex, and although it seems to be on the surface is in reality covered by the outer lamina of the cell. There is no regularity as to its place in the cell; it is not unfrequently, however, central or nearly so.

“As only one areola belongs to each cell, and as in many cases where it exists in the common cells of the epidermis, it is also visible in the cutaneous glands or stomata, and in these is always double–one being on each side of the limb–it is highly probable that the cutaneous gland is in all cases composed of two cells of peculiar form, the line of union being the longitudinal axis of the disk or pore.

“This areola, or nucleus of the cell as perhaps it might be termed, is not confined to the epidermis, being also found, not only in the pubescence of the surface, particularly when jointed, as in cypripedium, but in many cases in the parenchyma or internal cells of the tissue, especially when these are free from the deposition of granular matter.

“In the compressed cells of the epidermis the nucleus is in a corresponding degree flattened; but in the internal tissue it is often nearly spherical, more or less firmly adhering to one of the walls, and projecting into the cavity of the cell. In this state it may not unfrequently be found. in the substance of the column and in that of the perianthium.

“The nucleus is manifest also in the tissue of the stigma, where in accordance with the compression of the utriculi, it has an intermediate form, being neither so much flattened as in the epidermis nor so convex as it is in the internal tissue of the column.

“I may here remark that I am acquainted with one case of apparent exception to the nucleus being solitary in each utriculus or cell–namely, in Bletia Tankervilliae. In the utriculi of the stigma of this plant, I have generally, though not always, found a second areola apparently on the surface, and composed of much larger granules than the ordinary nucleus, which is formed of very minute granular matter, and seems to be deep seated.

“Mr. Bauer has represented the tissue of the stigma, in the species of Bletia, both before and, as he believes, after impregnation; and in the latter state the utriculi are marked with from one to three areolae of similar appearance.

“The nucleus may even be supposed to exist in the pollen of this family. In the early stages of its formation, at least a minute areola is of ten visible in the simple grain, and in each of the constituent parts of cells of the compound grain. But these areolae may perhaps rather be considered as merely the points of production of the tubes.

“This nucleus of the cell is not confined to orchideae, but is equally manifest in many other monocotyledonous families; and I have even found it, hitherto however in very few cases, in the epidermis of dicotyledonous plants; though in this primary division it may perhaps be said to exist in the early stages of development of the pollen. Among monocotyledons, the orders in which it is most remarkable are Liliaceae, Hemerocallideae, Asphodeleae, Irideae, and Commelineae.

“In some plants belonging to this last-mentioned family, especially in Tradascantia virginica, and several nearly related species, it is uncommonly distinct, not in the epidermis and in the jointed hairs of the filaments, but in the tissue of the stigma, in the cells of the ovulum even before impregnation, and in all the stages of formation of the grains of pollen, the evolution of which is so remarkable in tradascantia.

“The few indications of the presence of this nucleus, or areola, that I have hitherto met with in the publications of botanists are chiefly in some figures of epidermis, in the recent works of Meyen and Purkinje, and in one case, in M. Adolphe Broigniart’s memoir on the structure of leaves. But so little importance seems to be attached to it that the appearance is not always referred to in the explanations of the figures in which it is represented. Mr. Bauer, however, who has also figured it in the utriculi of the stigma of Bletia Tankervilliae has more particularly noticed it, and seems to consider it as only visible after impregnation.”[2]


That this newly recognized structure must be important in the economy of the cell was recognized by Brown himself, and by the celebrated German Meyen, who dealt with it in his work on vegetable physiology, published not long afterwards; but it remained for another German, the professor of botany in the University of Jena, Dr. M. J. Schleiden, to bring the nucleus to popular attention, and to assert its all-importance in the economy of the cell.

Schleiden freely acknowledged his indebtedness to Brown for first knowledge of the nucleus, but he soon carried his studies of that structure far beyond those of its discoverer. He came to believe that the nucleus is really the most important portion of the cell, in that it is the original structure from which the remainder of the cell is developed. Hence he named it the cytoblast. He outlined his views in an epochal paper published in Muller’s Archives in 1838, under title of “Beitrage zur Phytogenesis.” This paper is in itself of value, yet the most important outgrowth of Schleiden’s observations of the nucleus did not spring from his own labors, but from those of a friend to whom he mentioned his discoveries the year previous to their publication. This friend was Dr. Theodor Schwann, professor of physiology in the University of Louvain.

At the moment when these observations were communicated to him Schwann was puzzling over certain details of animal histology which he could not clearly explain. His great teacher, Johannes Muller, had called attention to the strange resemblance to vegetable cells shown by certain cells of the chorda dorsalis (the embryonic cord from which the spinal column is developed), and Schwann himself had discovered a corresponding similarity in the branchial cartilage of a tadpole. Then, too, the researches of Friedrich Henle had shown that the particles that make up the epidermis of animals are very cell-like in appearance. Indeed, the cell-like character of certain animal tissues had come to be matter of common note among students of minute anatomy. Schwann felt that this similarity could not be mere coincidence, but he had gained no clew to further insight until Schleiden called his attention to the nucleus. Then at once he reasoned that if there really is the correspondence between vegetable and animal tissues that he suspected, and if the nucleus is so important in the vegetable cell as Schleiden believed, the nucleus should also be found in the ultimate particles of animal tissues.

Schwann’s researches soon showed the entire correctness of this assumption. A closer study of animal tissues under the microscope showed, particularly in the case of embryonic tissues, that “opaque spots” such as Schleiden described are really to be found there in abundance–forming, indeed, a most characteristic phase of the structure. The location of these nuclei at comparatively regular intervals suggested that they are found in definite compartments of the tissue, as Schleiden had shown to be the case with vegetables; indeed, the walls that separated such cell-like compartments one from another were in some cases visible. Particularly was this found to be the case with embryonic tissues, and the study of these soon convinced Schwann that his original surmise had been correct, and that all animal tissues are in their incipiency composed of particles not unlike the ultimate particles of vegetables in short, of what the botanists termed cells. Adopting this name, Schwann propounded what soon became famous as his cell theory, under title of Mikroskopische Untersuchungen uber die Ubereinstimmung in der Structur und dent Wachsthum der Thiere und Pflanzen. So expeditious had been his work that this book was published early in 1839, only a few months after the appearance of Schleiden’s paper.

As the title suggests, the main idea that actuated Schwann was to unify vegetable and animal tissues. Accepting cell-structure as the basis of all vegetable tissues, he sought to show that the same is true of animal tissues, all the seeming diversities of fibre being but the alteration and development of what were originally simple cells. And by cell Schwann meant, as did Schleiden also, what the word ordinarily implies–a cavity walled in on all sides. He conceived that the ultimate constituents of all tissues were really such minute cavities, the most important part of which was the cell wall, with its associated nucleus. He knew, indeed, that the cell might be filled with fluid contents, but he regarded these as relatively subordinate in importance to the wall itself. This, however, did not apply to the nucleus, which was supposed to lie against the cell wall and in the beginning to generate it. Subsequently the wall might grow so rapidly as to dissociate itself from its contents, thus becoming a hollow bubble or true cell; but the nucleus, as long as it lasted, was supposed to continue in contact with the cell wall. Schleiden had even supposed the nucleus to be a constituent part of the wall, sometimes lying enclosed between two layers of its substance, and Schwann quoted this view with seeming approval. Schwann believed, however, that in the mature cell the nucleus ceased to be functional and disappeared.

The main thesis as to the similarity of development of vegetable and animal tissues and the cellular nature of the ultimate constitution of both was supported by a mass of carefully gathered evidence which a multitude of microscopists at once confirmed, so Schwann’s work became a classic almost from the moment of its publication. Of course various other workers at once disputed Schwann’s claim to priority of discovery, in particular the English microscopist Valentin, who asserted, not without some show of justice, that he was working closely along the same lines. Put so, for that matter, were numerous others, as Henle, Turpin, Du-mortier, Purkinje, and Muller, all of whom Schwann himself had quoted. Moreover, there were various physiologists who earlier than any of these had foreshadowed the cell theory–notably Kaspar Friedrich Wolff, towards the close of the previous century, and Treviranus about 1807, But, as we have seen in so many other departments of science, it is one thing to foreshadow a discovery, it is quite another to give it full expression and make it germinal of other discoveries. And when Schwann put forward the explicit claim that “there is one universal principle of development for the elementary parts, of organisms, however different, and this principle is the formation of cells,” he enunciated a doctrine which was for all practical purposes absolutely new and opened up a novel field for the microscopist to enter. A most important era in physiology dates from the publication of his book in 1839.


That Schwann should have gone to embryonic tissues for the establishment of his ideas was no doubt due very largely to the influence of the great Russian Karl Ernst von Baer, who about ten years earlier had published the first part of his celebrated work on embryology, and whose ideas were rapidly gaining ground, thanks largely to the advocacy of a few men, notably Johannes Muller, in Germany, and William B. Carpenter, in England, and to the fact that the improved microscope had made minute anatomy popular. Schwann’s researches made it plain that the best field for the study of the animal cell is here, and a host of explorers entered the field. The result of their observations was, in the main, to confirm the claims of Schwann as to the universal prevalence of the cell. The long-current idea that animal tissues grow only as a sort of deposit from the blood-vessels was now discarded, and the fact of so-called plantlike growth of animal cells, for which Schwann contended, was universally accepted. Yet the full measure of the affinity between the two classes of cells was not for some time generally apprehended.

Indeed, since the substance that composes the cell walls of plants is manifestly very different from the limiting membrane of the animal cell, it was natural, so long as the, wall was considered the most essential part of the structure, that the divergence between the two classes of cells should seem very pronounced. And for a time this was the conception of the matter that was uniformly accepted. But as time went on many observers had their attention called to the peculiar characteristics of the contents of the cell, and were led to ask themselves whether these might not be more important than had been supposed. In particular, Dr. Hugo von Mohl, professor of botany in the University of Tubingen, in the course of his exhaustive studies of the vegetable cell, was impressed with the peculiar and characteristic appearance of the cell contents. He observed universally within the cell “an opaque, viscid fluid, having granules intermingled in it,” which made up the main substance of the cell, and which particularly impressed him because under certain conditions it could be seen to be actively in motion, its parts separated into filamentous streams.

Von Mohl called attention to the fact that this motion of the cell contents had been observed as long ago as 1774 by Bonaventura Corti, and rediscovered in 1807 by Treviranus, and that these observers had described the phenomenon under the “most unsuitable name of ‘rotation of the cell sap.’ Von Mohl recognized that the streaming substance was something quite different from sap. He asserted that the nucleus of the cell lies within this substance and not attached to the cell wall as Schleiden had contended. He saw, too, that the chlorophyl granules, and all other of the cell contents, are incorporated with the “opaque, viscid fluid,” and in 1846 he had become so impressed with the importance of this universal cell substance that be gave it the name of protoplasm. Yet in so doing he had no intention of subordinating the cell wall. The fact that Payen, in 1844, had demonstrated that the cell walls of all vegetables, high or low, are composed largely of one substance, cellulose, tended to strengthen the position of the cell wall as the really essential structure, of which the protoplasmic contents were only subsidiary products.

Meantime, however, the students of animal histology were more and more impressed with the seeming preponderance of cell contents over cell walls in the tissues they studied. They, too, found the cell to be filled with a viscid, slimy fluid capable of motion. To this Dujardin gave the name of sarcode. Presently it came to be known, through the labors of Kolliker, Nageli, Bischoff, and various others, that there are numerous lower forms of animal life which seem to be composed of this sarcode, without any cell wall whatever. The same thing seemed to be true of certain cells of higher organisms, as the blood corpuscles. Particularly in the case of cells that change their shape markedly, moving about in consequence of the streaming of their sarcode, did it seem certain that no cell wall is present, or that, if present, its role must be insignificant.

And so histologists came to question whether, after all, the cell contents rather than the enclosing wall must not be the really essential structure, and the weight of increasing observations finally left no escape from the conclusion that such is really the case. But attention being thus focalized on the cell contents, it was at once apparent that there is a far closer similarity between the ultimate particles of vegetables and those of animals than had been supposed. Cellulose and animal membrane being now regarded as more by-products, the way was clear for the recognition of the fact that vegetable protoplasm and animal sarcode are marvellously similar in appearance and general properties. The closer the observation the more striking seemed this similarity; and finally, about 1860, it was demonstrated by Heinrich de Bary and by Max Schultze that the two are to all intents and purposes identical. Even earlier Remak had reached a similar conclusion, and applied Von Mohl’s word protoplasm to animal cell contents, and now this application soon became universal. Thenceforth this protoplasm was to assume the utmost importance in the physiological world, being recognized as the universal “physical basis of life,” vegetable and animal alike. This amounted to the logical extension and culmination of Schwann’s doctrine as to the similarity of development of the two animate kingdoms. Yet at the, same time it was in effect the banishment of the cell that Schwann had defined. The word cell was retained, it is true, but it no longer signified a minute cavity. It now implied, as Schultze defined it, “a small mass of protoplasm endowed with the attributes of life.” This definition was destined presently to meet with yet another modification, as we shall see; but the conception of the protoplasmic mass as the essential ultimate structure, which might or might not surround itself with a protective covering, was a permanent addition to physiological knowledge. The earlier idea had, in effect, declared the shell the most important part of the egg; this developed view assigned to the yolk its true position.

In one other important regard the theory of Schleiden and Schwann now became modified. This referred to the origin of the cell. Schwann had regarded cell growth as a kind of crystallization, beginning with the deposit of a nucleus about a granule in the intercellular substance–the cytoblastema, as Schleiden called it. But Von Mohl, as early as 1835, had called attention to the formation of new vegetable cells through the division of a pre-existing cell. Ehrenberg, another high authority of the time, contended that no such division occurs, and the matter was still in dispute when Schleiden came forward with his discovery of so-called free cell-formation within the parent cell, and this for a long time diverted attention from the process of division which Von Mohl had described. All manner of schemes of cell-formation were put forward during the ensuing years by a multitude of observers, and gained currency notwithstanding Von Mohl’s reiterated contention that there are really but two ways in which the formation of new cells takes place–namely, “first, through division of older cells; secondly, through the formation of secondary cells lying free in the cavity of a cell.”

But gradually the researches of such accurate observers as Unger, Nageli, Kolliker, Reichart, and Remak tended to confirm the opinion of Von Mohl that cells spring only from cells, and finally Rudolf Virchow brought the matter to demonstration about 1860. His Omnis cellula e cellula became from that time one of the accepted data of physiology. This was supplemented a little later by Fleming’s Omnis nucleus e nucleo, when still more refined methods of observation had shown that the part of the cell which always first undergoes change preparatory to new cell-formation is the all-essential nucleus. Thus the nucleus was restored to the important position which Schwann and Schleiden had given it, but with greatly altered significance. Instead of being a structure generated de novo from non-cellular substance, and disappearing as soon as its function of cell-formation was accomplished, the nucleus was now known as the central and permanent feature of every cell, indestructible while the cell lives, itself the division-product of a pre-existing nucleus, and the parent, by division of its substance, of other generations of nuclei. The word cell received a final definition as “a small mass of protoplasm supplied with a nucleus.”

In this widened and culminating general view of the cell theory it became clear that every animate organism, animal or vegetable, is but a cluster of nucleated cells, all of which, in each individual case, are the direct descendants of a single primordial cell of the ovum. In the developed individuals of higher organisms the successive generations of cells become marvellously diversified in form and in specific functions; there is a wonderful division of labor, special functions being chiefly relegated to definite groups of cells; but from first to last there is no function developed that is not present, in a primitive way, in every cell, however isolated; nor does the developed cell, however specialized, ever forget altogether any one of its primordial functions or capacities. All physiology, then, properly interpreted, becomes merely a study of cellular activities; and the development of the cell theory takes its place as the great central generalization in physiology of the nineteenth century. Something of the later developments of this theory we shall see in another connection.


Just at the time when the microscope was opening up the paths that were to lead to the wonderful cell theory, another novel line of interrogation of the living organism was being put forward by a different set of observers. Two great schools of physiological chemistry had arisen–one under guidance of Liebig and Wohler, in Germany, the other dominated by the great French master Jean Baptiste Dumas. Liebig had at one time contemplated the study of medicine, and Dumas had achieved distinction in connection with Prevost, at Geneva, in the field of pure physiology before he turned his attention especially to chemistry. Both these masters, therefore, and Wohler as well, found absorbing interest in those phases of chemistry that have to do with the functions of living tissues; and it was largely through their efforts and the labors of their followers that the prevalent idea that vital processes are dominated by unique laws was discarded and physiology was brought within the recognized province of the chemist. So at about the time when the microscope had taught that the cell is the really essential structure of the living organism, the chemists had come to understand that every function of the organism is really the expression of a chemical change–that each cell is, in short, a miniature chemical laboratory. And it was this combined point of view of anatomist and chemist, this union of hitherto dissociated forces, that made possible the inroads into the unexplored fields of physiology that were effected towards the middle of the nineteenth century.

One of the first subjects reinvestigated and brought to proximal solution was the long-mooted question of the digestion of foods. Spallanzani and Hunter had shown in the previous century that digestion is in some sort a solution of foods; but little advance was made upon their work until 1824, when Prout detected the presence of hydrochloric acid in the gastric juice. A decade later Sprott and Boyd detected the existence of peculiar glands in the gastric mucous membrane; and Cagniard la Tour and Schwann independently discovered that the really active principle of the gastric juice is a substance which was named pepsin, and which was shown by Schwann to be active in the presence of hydrochloric acid.

Almost coincidently, in 1836, it was discovered by Purkinje and Pappenheim that another organ than the stomach–namely, the pancreas–has a share in digestion, and in the course of the ensuing decade it came to be known, through the efforts of Eberle, Valentin, and Claude Bernard, that this organ is all-important in the digestion of starchy and fatty foods. It was found, too, that the liver and the intestinal glands have each an important share in the work of preparing foods for absorption, as also has the saliva–that, in short, a coalition of forces is necessary for the digestion of all ordinary foods taken into the stomach.

And the chemists soon discovered that in each one of the essential digestive juices there is at least one substance having certain resemblances to pepsin, though acting on different kinds of food. The point of resemblance between all these essential digestive agents is that each has the remarkable property of acting on relatively enormous quantities of the substance which it can digest without itself being destroyed or apparently even altered. In virtue of this strange property, pepsin and the allied substances were spoken of as ferments, but more recently it is customary to distinguish them from such organized ferments as yeast by designating them enzymes. The isolation of these enzymes, and an appreciation of their mode of action, mark a long step towards the solution of the riddle of digestion, but it must be added that we are still quite in the dark as to the real ultimate nature of their strange activity.

In a comprehensive view, the digestive organs, taken as a whole, are a gateway between the outside world and the more intimate cells of the organism. Another equally important gateway is furnished by the lungs, and here also there was much obscurity about the exact method of functioning at the time of the revival of physiological chemistry. That oxygen is consumed and carbonic acid given off during respiration the chemists of the age of Priestley and Lavoisier had indeed made clear, but the mistaken notion prevailed that it was in the lungs themselves that the important burning of fuel occurs, of which carbonic acid is a chief product. But now that attention had been called to the importance of the ultimate cell, this misconception could not long hold its ground, and as early as 1842 Liebig, in the course of his studies of animal heat, became convinced that it is not in the lungs, but in the ultimate tissues to which they are tributary, that the true consumption of fuel takes place. Reviving Lavoisier’s idea, with modifications and additions, Liebig contended, and in the face of opposition finally demonstrated, that the source of animal heat is really the consumption of the fuel taken in through the stomach and the lungs. He showed that all the activities of life are really the product of energy liberated solely through destructive processes, amounting, broadly speaking, to combustion occurring in the ultimate cells of the organism. Here is his argument:


“The oxygen taken into the system is taken out again in the same forms, whether in summer or in winter; hence we expire more carbon in cold weather, and when the barometer is high, than we do in warm weather; and we must consume more or less carbon in our food in the same proportion; in Sweden more than in Sicily; and in our more temperate climate a full eighth more in winter than in summer.

“Even when we consume equal weights of food in cold and warm countries, infinite wisdom has so arranged that the articles of food in different climates are most unequal in the proportion of carbon they contain. The fruits on which the natives of the South prefer to feed do not in the fresh state contain more than twelve per cent. of carbon, while the blubber and train-oil used by the inhabitants of the arctic regions contain from sixty-six to eighty per cent. of carbon.

“It is no difficult matter, in warm climates, to study moderation in eating, and men can bear hunger for a long time under the equator; but cold and hunger united very soon exhaust the body.

“The mutual action between the elements of the food and the oxygen conveyed by the circulation of the blood to every part of the body is the source of animal heat.

“All living creatures whose existence depends on the absorption of oxygen possess within themselves a source of heat independent of surrounding objects.

“This truth applies to all animals, and extends besides to the germination of seeds, to the flowering of plants, and to the maturation of fruits. It is only in those parts of the body to which arterial blood, and with it the oxygen absorbed in respiration, is conveyed that heat is produced. Hair, wool, or feathers do not possess an elevated temperature. This high temperature of the animal body, or, as it may be called, disengagement of heat, is uniformly and under all circumstances the result of the combination of combustible substance with oxygen.

“In whatever way carbon may combine with oxygen, the act of combination cannot take place without the disengagement of heat. It is a matter of indifference whether the combination takes place rapidly or slowly, at a high or at a low temperature; the amount of heat liberated is a constant quantity. The carbon of the food, which is converted into carbonic acid within the body, must give out exactly as much heat as if it had been directly burned in the air or in oxygen gas; the only difference is that the amount of heat produced is diffused over unequal times. In oxygen the combustion is more rapid and the heat more intense; in air it is slower, the temperature is not so high, but it continues longer.

“It is obvious that the amount of heat liberated must increase or diminish with the amount of oxygen introduced in equal times by respiration. Those animals which respire frequently, and consequently consume much oxygen, possess a higher temperature than others which, with a body of equal size to be heated, take into the system less oxygen. The temperature of a child (102 degrees) is higher than that of an adult (99.5 degrees). That of birds (104 to 105.4 degrees) is higher than that of quadrupeds (98.5 to 100.4 degrees), or than that of fishes or amphibia, whose proper temperature is from 3.7 to 2.6 degrees higher than that of the medium in which they live. All animals, strictly speaking, are warm-blooded; but in those only which possess lungs is the temperature of the body independent of the surrounding medium.

“The most trustworthy observations prove that in all climates, in the temperate zones as well as at the equator or the poles, the temperature of the body in man, and of what are commonly called warm-blooded animals, is invariably the same; yet how different are the circumstances in which they live.

“The animal body is a heated mass, which bears the same relation to surrounding objects as any other heated mass. It receives heat when the surrounding objects are hotter, it loses heat when they are colder than itself. We know that the rapidity of cooling increases with the difference between the heated body and that of the surrounding medium–that is, the colder the surrounding medium the shorter the time required for the cooling of the heated body. How unequal, then, must be the loss of heat of a man at Palermo, where the actual temperature is nearly equal to that of the body, and in the polar regions, where the external temperature is from 70 to 90 degrees lower.

“Yet notwithstanding this extremely unequal loss of heat, experience has shown that the blood of an inhabitant of the arctic circle has a temperature as high as that of the native of the South, who lives in so different a medium. This fact, when its true significance is perceived, proves that the heat given off to the surrounding medium is restored within the body with great rapidity. This compensation takes place more rapidly in winter than in summer, at the pole than at the equator.

“Now in different climates the quantity of oxygen introduced into the system of respiration, as has been already shown, varies according to the temperature of the external air; the quantity of inspired oxygen increases with the loss of heat by external cooling, and the quantity of carbon or hydrogen necessary to combine with this oxygen must be increased in like ratio. It is evident that the supply of heat lost by cooling is effected by the mutual action of the elements of the food and the inspired oxygen, which combine together. To make use of a familiar, but not on that account a less just illustration, the animal body acts, in this respect, as a furnace, which we supply with fuel. It signifies nothing what intermediate forms food may assume, what changes it may undergo in the body, the last change is uniformly the conversion of carbon into carbonic acid and of its hydrogen into water; the unassimilated nitrogen of the food, along with the unburned or unoxidized carbon, is expelled in the excretions. In order to keep up in a furnace a constant temperature, we must vary the supply of fuel according to the external temperature–that is, according to the supply of oxygen.

“In the animal body the food is the fuel; with a proper supply of oxygen we obtain the heat given out during its oxidation or combustion.”[3]


Further researches showed that the carriers of oxygen, from the time of its absorption in the lungs till its liberation in the ultimate tissues, are the red corpuscles, whose function had been supposed to be the mechanical one of mixing of the blood. It transpired that the red corpuscles are composed chiefly of a substance which Kuhne first isolated in crystalline form in 1865, and which was named haemoglobin–a substance which has a marvellous affinity for oxygen, seizing on it eagerly at the lungs vet giving it up with equal readiness when coursing among the remote cells of the body. When freighted with oxygen it becomes oxyhaemoglobin and is red in color; when freed from its oxygen it takes a purple hue; hence the widely different appearance of arterial and venous blood, which so puzzled the early physiologists.

This proof of the vitally important role played by the red-blood corpuscles led, naturally, to renewed studies of these infinitesimal bodies. It was found that they may vary greatly in number at different periods in the life of the same individual, proving that they may be both developed and destroyed in the adult organism. Indeed, extended observations left no reason to doubt that the process of corpuscle formation and destruction may be a perfectly normal one–that, in short, every red-blood corpuscle runs its course and dies like any more elaborate organism. They are formed constantly in the red marrow of bones, and are destroyed in the liver, where they contribute to the formation of the coloring matter of the bile. Whether there are other seats of such manufacture and destruction of the corpuscles is not yet fully determined. Nor are histologists agreed as to whether the red-blood corpuscles themselves are to be regarded as true cells, or merely as fragments of cells budded out from a true cell for a special purpose; but in either case there is not the slightest doubt that the chief function of the red corpuscle is to carry oxygen.

If the oxygen is taken to the ultimate cells before combining with the combustibles it is to consume, it goes without saying that these combustibles themselves must be carried there also. Nor could it be in doubt that the chiefest of these ultimate tissues, as regards, quantity of fuel required, are the muscles. A general and comprehensive view of the organism includes, then, digestive apparatus and lungs as the channels of fuel-supply; blood and lymph channels as the transportation system; and muscle cells, united into muscle fibres, as the consumption furnaces, where fuel is burned and energy transformed and rendered available for the purposes of the organism, supplemented by a set of excretory organs, through which the waste products–the ashes–are eliminated from the system.

But there remain, broadly speaking, two other sets of organs whose size demonstrates their importance in the economy of the organism, yet whose functions are not accounted for in this synopsis. These are those glandlike organs, such as the spleen, which have no ducts and produce no visible secretions, and the nervous mechanism, whose central organs are the brain and spinal cord. What offices do these sets of organs perform in the great labor-specializing aggregation of cells which we call a living organism?

As regards the ductless glands, the first clew to their function was given when the great Frenchman Claude Bernard (the man of whom his admirers loved to say, “He is not a physiologist merely; he is physiology itself”) discovered what is spoken of as the glycogenic function of the liver. The liver itself, indeed, is not a ductless organ, but the quantity of its biliary output seems utterly disproportionate to its enormous size, particularly when it is considered that in the case of the human species the liver contains normally about one-fifth of all the blood in the entire body. Bernard discovered that the blood undergoes a change of composition in passing through the liver. The liver cells (the peculiar forms of which had been described by Purkinje, Henle, and Dutrochet about 1838) have the power to convert certain of the substances that come to them into a starchlike compound called glycogen, and to store this substance away till it is needed by the organism. This capacity of the liver cells is quite independent of the bile-making power of the same cells; hence the discovery of this glycogenic function showed that an organ may have more than one pronounced and important specific function. But its chief importance was in giving a clew to those intermediate processes between digestion and final assimilation that are now known to be of such vital significance in the economy of the organism.

In the forty odd years that have elapsed since this pioneer observation of Bernard, numerous facts have come to light showing the extreme importance of such intermediate alterations of food-supplies in the blood as that performed by the liver. It has