What is Protoplasm?—Enough has now been given of disclosures of the modern microscope to show that our old friend Protoplasm has assumed an entirely new guise, if indeed it has not disappeared altogether. These simplest life processes are so marvelous and involve the action of such an intricate mass of machinery that we can no longer retain our earlier notion of protoplasm as the physical basis of life. There can be no life without the properties of assimilation, growth, and reproduction; and, so far as we know, these properties are found only in that combination of bodies which we call the cell, with its mixture of harmoniously acting parts. Life, at least the life of a cell, is then not the property of a chemical compound protoplasm, but is the result of the activities of a machine. Indeed, we are now at a loss to know how we can retain the term protoplasm. As originally used it meant the contents of the cell, and the significance in the term was in the conception of protoplasm as a somewhat homogeneous chemical compound uniform in all types of life. But we now see that this cell contains not a single substance, but a large number, including solids, jelly masses, and liquids, each of which has its own chemical composition. The number of chemical compounds existing in the material formerly called protoplasm no one knows, but we do know that they are many, and that the different substances are combined to form a physical structure. Which of these various bodies shall we continue to call protoplasm? Shall it be the linin, or the liquids, or the microsomes, or the chromatin threads, or the centrosomes? Which of these is the actual physical basis of life? From the description of cell life which we have given, it will be evident that no one of them is a material upon which our chemical biologists can longer found a chemical theory of life. That chemical theory of life, as we have seen, was founded upon the conception that the primitive life substance is a definite chemical compound. No such compound has been discovered, and these disclosures of the microscope of the last few years have been such as to lead us to abandon hope of ever discovering such a compound. It is apparently impossible to reduce life to any simpler basis than this combination of bodies which make up what was formerly called protoplasm. The term protoplasm is still in use with different meanings as used by different writers. Sometimes it is used to refer to the entire contents of the cell; sometimes to the cell substance only outside the nucleus. Plainly, it is not the protoplasm of earlier years.

With this conclusion one of our fundamental questions has been answered. We found in our first chapter that the general activities of animals and plants are easily reduced to the action of a machine, provided we had the fundamental vital powers residing in the parts of that machine. We then asked whether these fundamental properties were themselves those of a chemical compound or whether they were to be reduced to the action of still smaller machines. The first answer which biologists gave to this question was that assimilation, growth, and reproduction were the simple properties of a complex chemical compound. This answer was certainly incorrect. Life activities are exhibited by no chemical compound, but, so far as we know, only by the machine called the cell. Thus it is that we are again reduced to the problem of understanding the action of a machine. It may be well to pause here a moment to notice that this position very greatly increases the difficulties in the way of a solution of the life problem. If the physical basis of life had proved to be a chemical compound, the problem of its origin would have been a chemical one. Chemical forces exist in nature, and these forces are sufficient to explain the formation of any kind of chemical compound. The problem of the origin of the life substance would then have been simply to account for certain conditions which resulted in such chemical combination as would give rise to this physical basis of life. But now that the simplest substance manifesting the phenomena of life is found to be a machine, we can no longer find in chemical forces efficient causes for its formation. Chemical forces and chemical affinity can explain chemical compounds of any degree of complexity, but they cannot explain the formation of machines. Machines are the result of forces of an entirely different nature. Man can manufacture machines by taking chemical compounds and putting them together into such relations that their interaction will give certain results. Bits of iron and steel, for instance, are put together to form a locomotive, but the action of the locomotive depends, not upon the chemical forces which made the steel, but upon the relation of the bits of steel to each other in the machine. So far as we have had any experience, machines have been built under the guidance of intelligence which adapts the parts to each other. When therefore we find that the simplest life substance is a machine, we are forced to ask what forces exist in nature which can in a similar way build machines by the adjustment of parts to each other. But this topic belongs to the second part of our subject, and must be for the present postponed.

Reaction against the Cell Doctrine.—As the knowledge of cells which we have outlined was slowly acquired, the conception of the cell passed through various modifications. At first the cell wall was looked upon as the fundamental part, but this idea soon gave place to the belief that it was the protoplasm that was alive. Under the influence of this thought the cell doctrine developed into something like the following: The cell is simply a bit of protoplasm and is the unit of living matter. The bodies of all larger animals and plants are made up of great numbers of these units acting together, and the activities of the entire organism are simply the sum of the activities of its cells. The organism is thus simply the sum of the cells which compose it, and its activities the sum of the activities of the individual cells. As more facts were disclosed the idea changed slightly. The importance of the nucleus became more and more forcibly impressed upon microscopists, and this body came after a little into such prominence as to hide from view the more familiar protoplasm. The marvellous activities of the nucleus soon caused it to be regarded as the important part of the cell, while all the rest was secondary. The cell was now thought of as a bit of nuclear matter surrounded by secondary parts. The marvellous activities of the nucleus, and above all, the fact that the nucleus alone is handed down from one generation to the next in reproduction, all attested to its great importance and to the secondary importance of the rest of the cell.

This was the most extreme position of the cell doctrine. The cell was the unit of living action, and the higher animal or plant simply a colony of such units. An animal was simply an association together for mutual advantage of independent units, just as a city is an association of independent individuals. The organization of the animals was simply the result of the combination of many independent units. There was no activity of the organism as a whole, but only of its independent parts. Cell life was superior to organized life. Just as, in a city, the city government is a name given to the combined action of the individuals, so are the actions of organisms simply the combined action of their individual cells.

Against such an extreme position there has been in recent years a decided reaction, and to-day it is becoming more and more evident that such a position cannot be maintained. In the first place, it is becoming evident that the cell substance is not to be entirely obliterated by the importance of the nucleus. That the nucleus is a most important vital centre is clear enough, but it is equally clear that nucleus and cell substance must be together to constitute the life substance. The complicated structure of the cell substance, the decided activity shown by its fibres in the process of cell division, clearly enough indicate that it is a part of the cell which can not be neglected in the study of the life substance. Again the discovery of the centrosome as a distinct morphological element has still further added to the complexity of the life substance, and proved that neither nucleus nor cell substance can be regarded as the cell or as constituting life. It is true that we may not yet know the source of this centrosome. We do not know whether it is handed down from generation to generation like the nucleus, or whether it can be made anew out of the cell substance in the life of an ordinary cell. But this is not material to its recognition as an organ of importance in the cell activity. Thus the cell proves itself not to; be a bit of nuclear matter surrounded by secondary parts, but a community of several perhaps equally important interrelated members.

Another series of observations weakened the cell doctrine in an entirely different direction. It had been assumed that the body of the multicellular animal or plant was made of independent units. Microscopists of a few years ago began to suggest that the cells are in reality not separated from each other, but are all connected by protoplasmic fibres. In quite a number of different kinds of tissue it has been determined that fine threads of protoplasmic material lead from one cell to another in such a way that the cells are in vital connection. The claim has been made that there is thus a protoplasmic connection between all the cells of the body of the animal, and that thus the animal or plant, instead of consisting of a large number of separate independent cells, consists of one great mass of living matter which is aggregated into little centres, each commonly holding a nucleus. Such a conclusion is not yet demonstrated, nor is its significance very clear should it prove to be a fact; but it is plain that such suggestions quite decidedly modify the conception of the body as a community of independent cells.

There is yet another line of thought which is weakening this early conception of the cell doctrine. There is a growing conviction that the view of the organism, simply as the sum of the activities of the individual cells, is not a correct understanding of it. According to this extreme position, a living thing can have no organization until it appears as the result of cell multiplication. To take a concrete case, the egg of a starfish can not possess any organization corresponding to the starfish. The egg is a single cell, and the starfish a community of cells. The egg can, therefore, no more contain the organization of a starfish than a hunter in the backwoods can contain within himself the organization of a great metropolis. The descendants of individuals like the hunter may unite to form a city, and the descendants of the egg cell may, by combining, give rise to the starfish. But neither can the man contain within himself the organization of the city, nor the egg that of the starfish. It is, perhaps, true that such an extreme position of the cell doctrine has not been held by any one, but thoughts very closely approximating to this view have been held by the leading advocates of the cell doctrine, and have beyond question been the inspiration of the development of that doctrine.

But certainly no such conception of the significance of cell structure would longer be held. In spite of the fact that the egg is a single cell, it is impossible to avoid the belief that in some way it contains the starfish. We need not, of course, think of it as containing the structure of a starfish, but we are forced to conclude that in some way its structure is such that it contains the starfish potentially. The relation of its parts and the forces therein are such that, when placed under proper conditions, it develops into a starfish. Another egg placed under identical conditions will develop into a sea urchin, and another into an oyster. If these three eggs have the power of developing into three different animals under identical conditions, it is evident that they must have corresponding differences in spite of the fact that each is a single cell. Each must in some way contain its corresponding adult. In other words, the organization must be within the cells, and hence not simply produced by the associations of cells.

Over this subject there has been a deal of puzzling and not a little experimentation. The presence of some sort of organization in the egg is clear—but what is meant by this statement is not quite so clear. Is this adult organization in the whole egg or only in its nucleus, and especially in the chromosomes which, as we have seen, contain the hereditary traits? When the egg begins to divide does each of the first two cells still contain potentially the organization of the whole adult, or only one half of it? Is the development of the egg simply the unfolding of some structure already present; or is the structure constantly developing into more and more complicated conditions owing to the bringing of its parts into new relations? To answer these questions experimenters have been engaged in dividing developing eggs into pieces to determine what powers are still possessed by the fragments. The results of such experiments are as yet rather conflicting, but it is evident enough from them that we can no longer look upon the egg cell as a simple undifferentiated cell. In some way it already contains the characters of the adult, and when we remember that the characters of the adult which are to be developed from the egg are already determined, even to many minute details—such, for instance, as the inheritance of a congenital mark—it becomes evident that the egg is a body of extraordinary complexity. And yet the egg is nothing more than a single cell agreeing with other cells in all its general characters. It is clear, then, that we must look upon organization as something superior to cells and something existing within them, or at least within the egg cell, and controlling its development. We are forced to believe, further, that there may be as important differences between two cells as there are between two adult animals or plants. In some way there must be concealed within the two cells which constitute the egg of the starfish and the man differences which correspond to the differences between the starfish and the man. Organization, in other words, is superior to cell structure, and the cell itself is an organization of smaller units.

As the result of these various considerations there has been, in recent years, something of a reaction against the cell doctrine as formerly held. While the study of cells is still regarded as the key to the interpretation of life phenomena, biologists are seeing more and more clearly that they must look deeper than simple cell structure for their explanation of the life processes. While the study of cells has thrown an immense amount of light upon life, we seem hardly nearer the centre of the problem than we were before the beginning of the series of discoveries inaugurated by the formulation of the doctrine of protoplasm.

Fundamental Vital Activities as Located in Cells.—We are now in position to ask whether our knowledge of cells has aided us in finding an explanation of the fundamental vital actions to which, as we have seen, life processes are to be reduced. The four properties of irritability, contractibility, assimilation, and reproduction, belong to these vital units—the cells, and it is these properties which we are trying to trace to their source as a foundation of vital activity.

We may first ask whether we have any facts which indicate that any special parts of the cell are associated with any of these fundamental activities. The first fact that stands out clearly is that the nucleus is connected most intimately with the process of reproduction and especially with heredity. This has long been believed, but has now been clearly demonstrated by the experiments of cutting into fragments the cell bodies of unicellular animals. As already noticed, those pieces which possess a nucleus are able to continue their life and reproduce themselves, while those without a nucleus are incapable of reproduction. With greater force still is the fact shown by the process of fertilization of the egg. The egg is very large and the male reproductive cell is very small, and the amount of material which the offspring derives from its mother is very great compared with that which it derives from its father. But the child inherits equally from father and mother, and hence we must find the hereditary traits handed down in some element which the offspring obtains equally from father and mother. As we have seen (Figs. 34-44), the only element which answers this demand is the nucleus, and more particularly the chromosomes of the nucleus. Clearly enough, then, we must look upon the nucleus as the special agent in reproduction of cells.

Again, we have apparently conclusive evidence that the nucleus controls that part of the assimilative process which we have spoken of as the constructive processes. The metabolic processes of life are both constructive and destructive. By the former, the material taken into the cell in the form of food is built up into cell tissue, such as linin, microsomes, etc., and, by the latter, these products are to a greater or less extent broken to pieces again to liberate their energy, and thus give rise to the activities of the cell. If the destructive processes were to go on alone the organism might continue to manifest its life activities for a time until it had exhausted the products stored up in its body for such purposes, but it would die from the lack of more material for destruction. Life is not complete without both processes. Now, in the life of the cell we may apparently attribute the destructive processes to the cell substance and the constructive processes to the nucleus. In a cell which has been cut into fragments those pieces without a nucleus continue to show the ordinary activities of life for a time, but they do not live very long (Fig. 25). The fragment is unable to assimilate its food sufficiently to build up more material. So long as it still retains within itself a sufficiency of already formed tissue for its destructive metabolism, it can continue to move around actively and behave like a complete cell, but eventually it dies from starvation. On the other hand, those fragments which retain a piece of the nucleus, even though they have only a small portion of the cell substance, feed, assimilate, and grow; in other words, they carry on not only the destructive but also the constructive changes. Plainly, this means that the nucleus controls the constructive processes, although it does not necessarily mean that the cell substance has no share in these constructive processes. Without the nucleus the cell is unable to perform those processes, while it is able to carry on the destructive processes readily enough. The nucleus controls, though it may not entirely carry on, the constructive metabolism.

It is equally clear that the cell substance is the seat of most of the destructive processes which constitute vital action. The cell substance is irritable, and is endowed with the power of contractility. Cell fragments without nucleii are sensitive enough, and can move around as readily as normal cells. Moreover, the various fibres which surround the centrosomes in cell division and whose contractions and expansions, as we have seen, pull the chromosomes apart in cell division, are parts of the cell substance. All of these are the results of destructive metabolism, and we must, therefore, conclude that destructive processes are seated in the cell substance.

The centrosome is too problematical as yet for much comment. It appears to be a piece of the machinery for bringing about cell division, but beyond this it is not safe to make any statements.

In brief, then, the cell body is a machine for carrying on destructive chemical changes, and liberating from the compounds thus broken to pieces their inclosed energy, which is at once converted into motion or heat or some other form of active energy. This chemical destruction is, however, possible only after the chemical compounds have become a part of the cell. The cell, therefore, possesses a nucleus which has the power of enabling it to assimilate its food—that is, to convert it into its own substance. The nucleus further contains a marvellous material—chromatin—which in someway exercises a controlling influence in its life and is handed down from one generation to another by continuous descent. Lastly, the cell has the centrosome, which brings about cell division in such a manner that this chromatin material is divided equally among the subsequent descendants, and thus insures that the daughter cells shall all be equivalent to each other and to the mother cell.

We must therefore look upon the organic cell as a little engine with admirably adapted parts. Within this engine chemical activity is excited. The fuel supplied to the engine is combined by chemical forces with the oxygen of the air. The vigour of the oxidation is partly dependent upon temperature, just as it is in any other oxidation process, and is of course dependent upon the presence of fuel to be oxidized, and air to furnish the oxygen. Unless the fuel is supplied and the air has free access to it, the machine stops, the cell dies. The energy liberated in this machine is converted into motion or some other form. We do not indeed understand the construction of the machine well enough to explain the exact mechanism by which this conversion takes place, but that there is such a mechanism can not be doubted, and the structure of the cell is certainly complex enough to give plenty of room for it. The irritability of the cell is easily understood; for, since it is made of very unstable chemical compounds, any slight disturbance or stimulation on one part will tend to upset its chemical stability and produce reaction; and this is what is meant by irritability.

Or, again, we may look upon the cell as a little chemical laboratory, where chemical changes are constantly occurring. These changes we do not indeed understand, but they are undoubtedly chemical changes. The result is that some compounds are pulled to pieces and part of the fragments liberated or excreted, while other parts are retained and built into other more complex compounds. The compounds thus manufactured are retained in the cell body, and it grows in bulk. This continues until the cell becomes too big, and then it divides.

If a machine is broken it ceases to carry on its proper duties, and if the parts are badly broken it is ruined. So with the cell. If it is broken by any means, mechanical, thermal, or otherwise, it ceases to run—we say it dies. It has within itself great power of repairing injury, and therefore it does not cease to act until the injury is so great as to be beyond repair. Thus it only stops its motion when the machinery has become so badly injured as to be beyond hope of repair, and hence the cell, after once ceasing its action, can never resume it again.

There are, of course, other functions of living things besides the few simple ones which we have considered. But these are the fundamental ones; and if we can reduce them to an intelligible explanation, we may feel that we have really grasped the essence of life. If we understand how the cell can move and grow and reproduce itself, we may rest assured that the other phenomena of life follow as a natural consequence. If, therefore, we have obtained an understanding of these fundamental vital phenomena, we have accomplished our object of comprehending the life phenomena in our chemical and mechanical laws.

But have we thus reduced these fundamental phenomena to an intelligible explanation? It must be acknowledged that we have not. We have reduced them to the action of chemical forces acting in a machine. But the machine itself is unintelligible. The organic cell is no more intelligible to us than is the body as a whole. The chemical understanding which we thought we had a few years ago in protoplasm has failed us, and nothing has taken its place We have no conception of what may be the primitive life substance. All we can say is that this most marvellous of all natural phenomena occurs only within that peculiar piece of machinery which we call the cell, and that it is the result of the action of physical forces in that machine. How the machine acts, or even the structure of the machine, we are as far from understanding as we were fifty years ago. The solution has retreated before us even faster than we have advanced toward it.

Summary.—We may now notice in a brief summary the position which we have reached. In our attempt to explain the living organism on the principle of the machine, we are very successful so far as secondary problems are concerned. Digestion, circulation, respiration, and motion are readily solved upon chemical and mechanical principles. Even the phenomena of the nervous system are, in a measure, capable of comprehension within a mechanical formula, leaving out of account the purely mental phenomena which certainly have not been touched by the investigation. All of these phenomena are reducible to a few simple fundamental activities, and these fundamental activities we find manifested by simple bits of living matter unincumbered by the complicated machinery of organisms. With the few fundamental properties of these bits of organic matter we can construct the complicated life of the higher organism. When we come, however, to study these simple bits of matter, they prove to be anything but simple bits of matter. They, too, are pieces of complicated mechanism whose action we do not even hope to understand. That their action is dependent upon their machinery is evident enough from the simple description of cell activity which we have noticed. That these fundamental vital properties are to be explained as the result of chemical and mechanical forces acting through this machinery, can not be doubted. But how this occurs or what constitutes the guiding force which corresponds to the engineer of the machine, we do not know.

Thus our mechanical explanation of the living machine lacks a foundation. We can understand tolerably well the building of the superstructure, but the foundation stones upon which that structure is built are unintelligible to us. The running of the living machine is thus only in part understood. The living organism is a machine or, it is better to say, it is a series of machines one within the other. As a whole it is a machine, and its parts are separate machines. Each part is further made up of still smaller machines until we reach the realm of the microscope. Here still we find the same story. Even the parts formerly called units, prove to be machines, and when we recognize the complexity of these cells and their marvellous activities, we are ready to believe that we may find still further machines within. And thus vital activity is reduced to a complex of machines, all acting in harmony with each other to produce together the one result—life.



PART II.

THE BUILDING OF THE LIVING MACHINE.

* * * * *

CHAPTER III.

THE FACTORS CONCERNED IN THE BUILDING OF THE LIVING MACHINE.

Having now outlined the results of our study into the mechanism of the living machine, we turn our attention next to the more difficult problem of the method by which this machine was built. From the facts which we have been considering in the last two chapters it is evident that the problem we have before us is a mechanical rather than a chemical one. Of course, chemical forces lie at the bottom of vital activity, and we must look upon the force of chemical affinity as the fundamental power to which the problems must be referred. But a chemical explanation will evidently not suffice for our purpose; for we have absolutely no reason for believing that the phenomena of life can occur as the results of the chemical properties of any compound, however complex. The simplest known form of matter which manifests life is a machine, and the problem of the origin of life must be of the origin of that machine. Are there any forces in nature which are of a sort as to enable us to use them to explain the building of machines? Plants and animals are the only machines which nature has produced. They are the only instances in nature of a structure built with its parts harmoniously adjusted to each other to the performance of certain ends. All other machines with which we are acquainted were made by man, and in making them intelligence came in to adapt the parts to each other. But in the living organism is a similarly adapted machine made by natural means rather than artificial. How were they built? Does nature, apart from human intelligence, possess forces which can achieve such results?

Here again we must attack the problem from what seems to be the wrong end. Apparently it would be simpler to discover the method of the manufacture of the simplest machine rather than the more complex ones. But this has proved contrary to the fact. Perhaps the chief reason is that the simplest living machine is the cell whose study must always involve the use of the microscope, and for this reason is more difficult. Perhaps it is because the problem is really a more difficult one than to explain the building of the more complex machines out of the simpler ones. At all events, the last fifty years have told us much of the method of the building of the complex machines out of the simpler ones, while we have as yet not even a hint as to the solution of the building of the simplest machine from the inanimate world. Our attention must, therefore, be first directed to the method by which nature has constructed the complex machines which we find filling the world to-day in the form of animals and plants.

History of the Living Machine.—In the first place, we must notice that these machines have not been fashioned suddenly or rapidly, but have been the result of a very slow growth. They have had a history extending very far back into the past for a period of years which we can only indefinitely estimate, but certainly reaching into the millions. As we look over this past history in the light of our present knowledge we see that whatever have been the forces which have been concerned in the construction of these machines they have acted very slowly. It has taken centuries, and, indeed, thousands of years, to take the successive steps which have been necessary in this construction. Secondly, we notice that the machines have been built up step by step, one feature being added to another with the slowly progressing ages. Thirdly, we notice that in one respect this construction of the living machine by nature's processes has been different from our ordinary method of building machines. Our method of building puts the parts gradually into place in such a way that until the machine is finished it is incapable of performing its functions. The half-built engine is as useless and as powerless as so much crude iron. Its power of action only appears after the last part is fitted into place and the machine finished. But nature's process in machine building is different. Every step in the process, so far as we can trace it at least, has produced a complete machine. So far back as we can follow this history we find that at every point the machine was so complete as to be always endowed with motion and life activity. Nature's method has been to take simpler types of machines and slowly change them into more complicated ones without at any moment impairing their vigour. It is something as if the steam engine of Watt should be slowly changed by adding piece after piece until there was finally produced the modern quadruple expansion engine, but all this change being made upon the original engine without once stopping its motion.



This gradual construction of the living machines has been called Organic Evolution, or the Theory of Descent. It will be necessary for us, in order to comprehend the problem which we have before us, to briefly outline the course of this evolution. Our starting point in this history must be the cell, for such is the earliest and simplest form of living thing of which we have any trace. This cell is, of course, already a machine, and we must presently return to the problem of its origin. At present we will assume this cell as a starting point endowed with its fundamental vital powers. It was sensitive, it could feel, grow, and reproduce itself. From such a simple machine, thus endowed, the history has been something as follows: In reproducing itself this machine, as we have already seen, simply divided itself into two halves, each like the other. At first all the parts thus arising separated from each other and remained independent. But so long as this habit continued there could be little advance. After a time some of the cells failed to separate after division, but remained clinging together (Fig. 45). The cells of such a mass must have been at first all alike; but, after a little, differences began to appear among them. Those on the outside of the mass were differently affected by their surroundings from those in the interior, and soon the cells began to share among themselves the different duties of life. The cells on the outside were better situated for protection and capturing food, while those on the inside could not readily seize food for themselves, and took upon themselves the duty of digesting the food which was handed to them by the outer cells. Each of these sets of cells could now carry on its own special duties to better advantage, since it was freed from other duties, and thus the whole mass of cells was better served than when each cell tried to do everything for itself. This was the first step in the building of the machine out of the active cells (Fig. 46). From such a starting point the subsequent history has been ever based upon the same principle. There has been a constant separation of the different functions of life among groups of cells, and as the history went on this division of labor among the different parts became greater and greater. Group after group of cells were set apart for one special duty after another, and the result was a larger and ever more complicated mass of cells, with a greater and greater differentiation among them. In this building of the machine there was no time when the machine was not active. At all points the machine was alive and functional, but each step made the total function of the machine a little more accurately performed, and hence raised somewhat the totality of life powers. This parcelling out of the different duties of life to groups of cells continued age after age, each step being a little advance over the last, until the result has been the living machine as we know it in its highest form, with its numerous organs, all interrelated in such a way as to form a harmoniously acting whole.



But a second principle in this growth of the machine was needed to produce the variety which is found in nature. As the different cells in the multicellular mass became associated into groups for different duties, the method of such division of labor was not alike in all machines. A city in China and one in America are alike made up of individuals, and the fundamental needs of the Chinaman and the American are alike. But differences in industrial and political conditions have produced different combinations and associations, so that Pekin is wonderfully unlike New York. So in these early developing machines, quite a variety of method of organization was adopted by the different groups. Now as soon as any special type of organization was adopted by any animal or plant, the principle of heredity transmitted the same kind of organization to its descendants, and there thus arose lines of descent differing from each other, each line having its own method of organization. As we follow the history of each line the same thing is repeated. We find that the representatives of each line again separate into groups, each of which has acquired some new type of organization, and there has thus been a constant divergence of these lines of descent in an indefinite number of directions. The members of the different lines of descent all show a fundamental likeness with each other since they retain the fundamental characters of their common ancestor, but they show also the differences which they have themselves acquired. And thus the process is repeated over and over again. This history of the growth of these different machines has thus been one of divergence from common centres, and is to be diagrammatically expressed after the fashion of a branching tree. The end of each branch represents the highest state of perfection to which each line has been carried.

One other point in this history must be noted. As the development of the complication of the machine progressed the possibility of further progress has been constantly narrowed. When the history of these machines began as a simple mass of cells, there was a possibility of an almost endless variety of methods of organization. But as a distinct type of organization was adopted by one and another line of descendants all subsequent productions were limited through the law of heredity to the general line of organization adopted by their ancestors. With each age the further growth of such machines must consist in the further development in the perfection of its parts, and not in the adoption of any new system of organization. Hence it is that the history of the living machine has shown a tendency toward development along a few well-marked lines, and although this complication becomes greater, we still see the same fundamental scheme of organization running through the whole. As the ages have progressed the machines have become more perfect in the adjustment of their parts, i.e., they have become more perfect machines, but the history has been simply that of perfecting the early machines rather than the production of new types.

Evidence for this History.—As just outlined, we see that the living machines have been gradually brought into their present condition by a process which has been called organic evolution. But we must pause for a moment to ask what is our evidence that such has been the history of the living machine. The whole possibility of understanding living nature depends upon our accepting this history and finding an explanation of it. At the outset we have the question of fact, and we must notice the grounds upon which we stand in assuming this history to be as outlined.

This problem is the one which has occupied such a prominent place in the scientific world during the last forty years, and which has contributed so largely toward making modern biology such a different subject from the earlier studies of natural history. It is simply the evidence for organic evolution, or the theory of descent. The subject has for forty years been thoroughly sifted and tested by every conceivable sort of test. As a result of the interest in the question there has been disclosed an immense mass of evidence, relevant and irrelevant. As the evidence has accumulated it has become more and more evident that the evolution theory must be recognized as the only one which is in accord with the facts, and the outcome has been a practical unanimity among thinkers that the theory of descent must be the foundation of our further study. The evidence which has forced this conclusion upon scientists we must stop for a moment to consider, since it bears very directly upon the subject we are studying.

Historical.—The first source of evidence is naturally a historical one. This long history of the construction of the living machine has left its record in the rocks which form the earth's surface. During this long period the rocks of the earth's crust have been deposited, and in these rocks have been left samples of many of the steps in this history of machine building. The history can be traced by the study of these samples just as the history of any machine might be traced from a study of the models in a patent office. One might very easily trace, with most strict accuracy and minute detail, the history of the printing machine from the models which are preserved in the patent offices and elsewhere. So is it with the history of the living machine. To be sure, the history is rather incomplete and at times difficult to read. Many a period in the development has left no samples for our inspection and must be interpreted in our history between what went before and what comes after. Many of the machines, especially the early ones, were made of such fragile material that they could not be preserved in the rocks. In many a case, too, the rocks in which the specimens were deposited have been subjected to such a variety of heatings and pressures, that they have been twisted out of shape and even crushed out of recognizable form. But in spite of this the record is showing itself more complete each year. Our paleontologists are opening layer after layer of these rocks, and thus examining each year new pages in nature's history. The more recent epochs in the history have been already read with almost historic accuracy. From them we have learned in great detail how the finishing touches were given to these machines, and are able to trace with accuracy how the somewhat more generalized forms of earlier days were changed to produce our modern animals.

This fossil record has given us our best knowledge of the course by which the present living world has been brought into its existing condition. But its accuracy is largely confined to the recent periods. Of the very early history fossils tell us little or nothing. All the early rocks, which we may believe were formed during the period when the first steps in this machine building were taken, have been so changed by heat and pressure that whatever specimens they may have originally contained have been crushed out of shape. Furthermore, the earliest organisms had no hard skeletons, and it was not until living beings had developed far enough to have hard parts that it was possible for them to leave traces of themselves in the rocks. Hence, so far as concerns this earliest history, we can get no record of it in the rocks.

Embryological.—But here comes in another source of evidence which helps to fill up the gap. In its development every animal to-day begins as an egg. This is a simple cell, and the animal goes through a series of changes which eventually lead to the adult. Now these changes appear for the most part to be parallel to the changes through which the earlier forms of life passed in their development from the simple to the more complicated forms. Where it is possible to follow the history of the groups of animals from their fossil remains and compare it with the history of the individual animal as it progresses from the egg to the adult, there is found a very decided parallelism. This parallelism between embryology and past history has been of great service in helping us toward the history of the past. At one time it was believed that it was the key which would unlock all doors, and for a decade biologists eagerly pursued embryology with the expectation that it would solve all problems in connection with the history of animals. The result has been somewhat disappointing. Embryology has, it is true, been of the utmost service in showing relationships of forms to each other, and in thus revealing past history. But while this record is a valuable one, it is a record which has unfortunately been subject to such modifying conditions that in many cases its original meaning has been entirely obliterated and it has become worthless as a historical record. These imperfections in regard to the record were early seen after the attention of biologists was seriously turned to the study of embryology, but it was expected that it would be possible to correct them and discover the true meaning underlying the more apparent one. Indeed, in many cases this has been found possible. But many of the modifications are so profound as to render it impossible to untangle them and discover the true meaning. As a result the biologist to-day is showing less confidence in embryology, and is turning his attention in different directions as more promising of results in the line desired.

But although the teachings of embryology have failed to realize the great hopes that were placed upon them, their assistance in the formulation of this history of the machine has been of extreme value. Many a bit of obscurity has been cleared up when the embryology of puzzling animals has been studied. Many a relationship has been made clear, and this is simply another way of saying that a portion of this history of life has been read. This aid of embryology has been particularly valuable in just that part of the history where the evidence from the study of fossils is wanting. The study of fossils, as we have seen, gives little or no data concerning the early history of living machines; and it is just here that embryology has proved to be of the most value. It is a source of evidence that has told us of most of the steps in the progress from the single-celled animal to the multicellular organisms, and gives us the clearest idea of the fundamental principles which have been concerned in the evolution of life and the construction of the complicated machine out of the simple bit of protoplasm. In spite of its limits, therefore, embryology has contributed a large quota of the evidence which we have of the evolution of life.

Anatomical.—A third source of this history is obtained from the facts of comparative anatomy. The essential feature of this subject is the fact that animals and plants show relationships. This fact is one of the most patent and yet one of the most suggestive facts of biology. It has been recognized from the very beginning of the study of animals and plants. One cannot be even the most superficial observer without seeing that certain forms show great likeness to each other while others are much more unlike. The grouping of animals and plants into orders, genera, and species is dependent upon this relationship. If two forms are alike in everything except some slight detail, they are commonly placed in the same genus but in different species, while if they show a greater unlikeness they may be placed in separate genera. By thus grouping together forms according to their resemblance the animal and vegetable kingdoms are classified into groups subordinate to groups. The principle of relationship, i.e., fundamental similarity of structure, runs through the whole animal and vegetable kingdom. Even the animals most unlike each other show certain points of similarity which indicates a relationship, although of course a distant one.

The fact of such a relationship is too patent to demand more words, but its significance needs to be pointed out. When we speak of relationship among men we always mean historical connection. Two brothers are closely related because they have sprung from common parents, while two cousins are less closely related because their common point of origin was farther back in time. More widely we speak of the relationship of the Indo-European races, meaning thereby that back in the history of man these races had a common point of origin. We never speak of any real relation of objects unless thereby we mean to imply historical connection. We are therefore justified in interpreting the manifest relationships of organisms as pointing to history. Particularly are we justified in this conclusion when we find that the relationships which we draw between the types of life now in existence run parallel to the history of these types as revealed to us by fossils and at the same time disclosed by the study of embryology.

This subject of comparative anatomy includes a consideration of what is called homology, and perhaps a concrete example may be instructive both in illustration and as suggesting the course which nature adopts in constructing her machines. We speak of a monkey's arm and a bird's wing as homologous, although they are wonderfully different in appearance and adapted to different duties. They are called homologous because they have similar parts in similar relations. This can be seen in Figs. 47 and 48, where it will be seen that each has the same bones, although in the bird's wing some of the bones have been fused together and others lost. Their similarity points to a relationship, but their dissimilarity tells us that the relationship is a distant one, and that their common point of origin must have been quite far back in history. Now if we follow back the history of these two kinds of appendages, as shown to us by fossils, we find them approaching a common point. The arm can readily be traced to a walking appendage, while the bird's wing, by means of some interesting connecting links, can in a similar way be traced to an appendage with its five fingers all free and used for walking. Fig. 49 shows one of these connecting links representing the earliest type of bird, where the fingers and bones of the arm were still distinct, and yet the whole formed a true wing. Thus we see that the common point of origin which is suggested by the likenesses between an arm and a wing is no mere imaginary one, for the fossil record has shown us the path leading to that point of origin. The whole tells us further that nature's method of producing a grasping or flying organ was here, not to build a new organ, but to take one that had hitherto been used for other purposes, and by slow changes modify its form and function until it was adapted to new duties.



Significance of these Sources of History.—The real force of these sources of evidence comes to us only when we compare them with each other. They agree in a most remarkable fashion. The history as disclosed by fossils and that told by embryology agree with each other, and these are in close harmony with the history as it can be read from comparative anatomy. If archaeologists were to find, in different countries and entirely unconnected with each other two or more different records of a lost nation, the belief in the actual existence of that nation would be irresistible. When researches at Nineveh, for example, unearth tablets which give the history of ancient nations, and when it proves that among the nations thus mentioned are some with the same names and having the same facts of history as those mentioned in the Bible, it is absolutely impossible to avoid the conclusion that such a nation with such a history did actually exist. Two independent sources of record could not be false in regard to such a matter as this.

Now, our sources of evidence for this history of the living machine prove to be of exactly this kind. We have three independent sources of evidence which are so entirely different from each other that there is almost no likeness between them. One is written in the rocks, one in bone and muscle, while the third is recorded in the evanescent and changing pages of embryology and metamorphosis. Yet each tells the same story. Each tells of a history of this machine from simple forms to more complex. Each tells of its greater and greater differentiation of labour and structure as the periods of time passed. Each tells of a growing complexity and an increasing perfection of the organisms as successive periods pass. Each tells us of common points of origin and divergence from these points. Each tells us how the more complicated forms have arisen as the results of changes in and modifications of the simpler forms. Each shows us how the individual parts of the organisms have been enlarged or diminished or changed in shape to adapt them to new duties. Each, in short, tells the same story of the gradual construction of the living machine by slow steps and through long ages of time. When these three sources of history so accurately agree with each other, it is as impossible to disbelieve in the existence of such history as it is to disbelieve in the existence of the ancient Hittite nation, after its history has been told to us by two different sources of record.

Now all this is very germane to our subject. We are trying to learn how this living machine, with its wonderful capabilities, was built. The history which we have outlined is undoubtedly the history of the building of this machine, and the knowledge that these complicated machines have been produced as the result of slow growth is of the utmost importance to us. This knowledge gives us at the very start some idea of the nature of the forces which have been at work. It tells us that in searching for these forces we must look for those which have been acting constantly. We must look for forces which produce their effects not by sudden additions to the complication of the machine. They must be constant forces whose effect at any one time is comparatively slight, but whose total effect is to increase the complexity of the machine. They must be forces which produce new types through the modification of the old ones. We must look for forces which do not adapt the machine for its future, but only for its present need. Each step in the history has been a complete animal with its own fully developed powers. We are not to expect to find forces which planned the perfect machine from the start, nor forces which were engaged in constructing parts for future use. Each step in the building of the machine was taken for the good of the machine at the particular moment, and the forces which we are to look for must therefore be only such as can adapt the organisms for its present needs. In other words, nothing has been produced in this machine for the purpose of being developed later into something of value, but all parts that have been produced are of value at the time of their appearance. We must, in short, look for forces constantly in action and always tending in the same direction of greater complexity of structure.

Is it possible to discover these forces and comprehend their action? Before the modern development of evolution this question would unhesitatingly have been answered in the negative. To-day, under the influence of the descent theory, stimulated, in the first place, by Darwin, the question will be answered by many with equal promptness in the affirmative. At all events, we have learned in the last forty years to recognize some of the factors which have been at work in the construction of this machine. We must turn, therefore, to the consideration of these factors.

Forces at Work in the Building of the Living Machine.—There are three primary factors which lie at the bottom of the whole process. They are—

1. Reproduction, which preserves type from generation to generation.

2. Variation, which modifies type from generation to generation.

3. Heredity, which transmits characters from generation to generation.

Each must be considered by itself.

Reproduction.—Reproduction is the primary factor in this process of machine building, heredity and variation being simply phases of reproduction. The living machine has developed by natural processes, all other machines by artificial methods. Reproduction is the one essential point of difference between the living machine and the others which has made their construction by natural processes a possibility. What, then, is reproduction? Reproduction is in all cases at the bottom simple division. Whether we consider the plant that multiplies by buds or the unicellular animal that simply divides into two equal parts, or the larger animal that multiplies by eggs, we find that in all cases the fundamental feature of the process is division. In all cases the organism divides into two or more parts, each of which becomes in time like the original. Moreover, when we trace this division further we find that in all cases it is to be referred back to the division of the cell, such as we have described in a previous chapter. The egg is a single cell which has come from the parent by the division of one of the cells in the body of the parent. A bud is simply a mass of cells which have all arisen from the parent cells by division. The foundation of reproduction is thus in all cases cell division. Now, this process of division is dependent upon the properties of the cell. Firstly, it is a result of the assimilative powers of the cell, for only through assimilation can the cell increase in size, and only as it increases in size can it gain sustenance for cell division. Secondly, it is dependent, as we have seen, upon the mechanism of the cell body, and especially the nucleus and centrosome. These structures regulate the cell division, and hence the reproduction of all animals and plants. We can not, therefore, find any explanation of reproduction until we have explained the mechanism of the cell. The fundamental feature, of nature's machine building is thus based upon the machinery of the nucleus and centrosome of the organic cell.

Aside from the simple fact that it preserves the race, the most important feature connected with this reproduction is its wonderful fruitfulness. Since it results from division, it always tends to increase the offspring in geometrical ratio. In the simplest case, that of the unicellular animals, the cell divides, giving rise to two animals, each of which divides again, producing four, and these again, giving eight, etc. The rapidity of this multiplication is sometimes inconceivable. It depends, of course, upon the interval of time between the successive divisions, but among the lower organisms this interval is sometimes not more than half an hour, the result of which is that a single individual could give rise in the course of twenty-four hours to sixteen million offspring. This is doubtless an extreme case, but among all the lower animals the rate is very great. Among larger animals the process is more complicated; but here, too, there is the same tendency to geometrical progression, although the intervals between the successive reproductions may be quite long and irregular. But it is always so great that if allowed to progress unhindered at its normal rate the offspring would, in a few years, become so numerous as to crowd other life out of existence. Even the slow-breeding elephant would, if allowed to breed unhindered for seven hundred and fifty years, produce nineteen million offspring—a rate of increase plainly incompatible with the continued existence of other animals.

Here, then, we have the foundation of nature's method of building animals and plants of the higher classes. In the machinery of the cell she has a power of reproduction which produces an increase in geometrical ratio far beyond the possibility for the surface of the earth to maintain.

Heredity.—The offspring which arise by these processes of division are like each other, and like the parent from which they sprung. This is the essence of what is called heredity. Its significance in the process of machine building is evident at once. It is the conserving force which preserves the forms already produced and makes it possible for each generation to build upon the structures of the earlier ones. Without it each generation would have to begin anew at the beginning, and nothing could be accomplished. But since this principle brings each individual to the same place where its parents stand, and thus always builds the offspring into a machine like the parent, it makes it possible for the successive generations to advance. Heredity is thus like the power of memory, or better still, like the invention of printing in the development of civilization. It is a record of past achievements. By means of printing each age is enabled to benefit by the discoveries of the previous age, and without it the development of civilization would be impossible. In the same way heredity enables each generation to benefit by the achievements of its ancestors in the process of machine building, and thus to devote its own energies to advancement.

The fact of heredity is patent enough. It has been always clearly recognized that the child has the characters of its parents, and this belief is so well attested as to need no proof. It is still a question as to just what characters may be inherited, and what influences may affect the inheritance. There are plenty of puzzling problems connected with heredity, but the fact of heredity is one of the foundation stones of biological science. Upon it must be built all theories which look toward the explanation of the origin of the living machine.

This factor of heredity again we must trace back to the machinery of the cell. We have seen in the previous pages evidence for the wonderful nature of the chromosomes of the cells. We can not pretend to understand them, but they must be extraordinarily complex. We have seen proof that these chromosomes are probably the physical basis of heredity, since they are the only parts of each parent which are handed down to subsequent generations. With these various facts of cell division and cell fertilization in mind, we can reach a very simple explanation of fundamental features of heredity. The following is an outline of the most widely accepted view of the hereditary process.

Recognizing that the chromosomes are the physical basis of hereditary transmission, we can picture to ourselves the transmission of hereditary characters something as follows: As we have seen, the fertilized egg contains an equal number of chromosomes from each parent (Fig. 42). Now when this fertilized cell divides, each of the rods splits lengthwise, half of each entering each of the two cells arising from the cell division. From this method of division of the chromosomes it follows that the daughter cells would be equivalent to each other and equivalent also to the undivided egg. If the original chromosomes contained potentially all the hereditary traits handed down from parent to child, the chromosomes of each daughter cell will contain similar hereditary traits. If, therefore, the original fertilized egg possessed the power of developing into an adult like the parent, each of the daughter cells should likewise possess the power of developing into a similar adult. And thus each cell which arises as the result of such division should possess similar characters so long as this method of division continues. But after a little in the development of the egg a differentiation among the daughter cells arises. They begin to acquire different shapes and different functions. This we can only believe to be the result of a differentiation in their chromatin material. In the cell division the chromosomes no longer split into equivalent halves, but some characters are portioned off to some cells and others to other cells. Those cells which are to carry on digestive functions when they are formed receive chromatin material which especially controls them in the performance of this digestive function, while those which are to produce sensory organs receive a different portion of the chromatin material. Thus the adult individual is built up as the cells receive different portions of this hereditary substance contained in the original chromosomes. The original chromosomes contained all hereditary characters, but as development proceeds these are gradually portioned out among the daughter cells until the adult is formed.

From this method of division it will be seen that each cell of the adult does not contain all the characters concealed in the original chromosomes of the egg, although each contains a part which may have been derived from each parent. It is thought, however, that a part of the original chromatin material does not thus become differentiated, but remains entirely unchanged as the individual is developing. This chromatin material may increase in amount by assimilation, but it remains unchanged during the entire growth of the individual. It thus follows that the adult will contain, along with its differentiated material, a certain amount of the original physical basis of heredity which still retains its original powers. This undifferentiated chromatin material originally possessed powers of producing a new individual, and of course it still possesses these powers, since it has remained dormant without alteration. Further, it will follow that if this dormant undifferentiated chromatin should start into activity and produce a new individual, the new individual thus produced would be identical in all characters with the one which actually did develop from the egg, since both individuals would have come from a bit of the same chromatin. The child would be like the parent. This would be true no matter how much this undifferentiated material should increase in amount by assimilation, so long as it remained unaltered in character, and it hence follows that every individual carries around a certain amount of undifferentiated chromatin material in all respects identical with that from which he developed.

Now whether this undifferentiated germ plasm, as we will now call it, is distributed all over the body, or is collected at certain points, is immaterial to our purpose. It is certain that portions of it find their way into the reproductive organs of the animal or plant. Thus we see that part of the chromatin material in the egg of the first generation develops into the second generation, while another part of it remains dormant in that second generation, eventually becoming the chromatin of its eggs and spermatozoa. Thus each egg of the second generation receives chromosomes which have come directly from the first generation, and thus it will follow that each of these eggs will have identical properties with the egg of the first generation. Hence if one of these new eggs develops into an adult it will produce an adult exactly like the second generation, since it contains chromosomes which are absolutely identical with those from which the second generation sprung. There is thus no difficulty in understanding why the second generation will be like the first, and since the process is simply repeated again in the next reproduction, the third generation will be like the second, and so on, generation after generation. A study of the accompanying diagram will make this clear.

In other words, we have here a simple understanding of at least some of the features of heredity. This explanation is that some of the chromatin material or germ plasm is handed down from one generation to another, and is stored temporarily in the nucleii of the reproductive cells. During the life of the individual this germ plasm is capable of increasing in amount without changing its nature, and it thus continues to grow and is handed down from generation to generation, always endowed with the power of developing into a new individual under proper conditions, and of course when it does thus give rise to new individuals they will all be alike. We can thus easily understand why a child is like its parent. It is not because the child can inherit directly from its parent, but rather because both child and parent have come from the unfolding of two bits of the same germ plasm. This fact of the transmission of the hereditary substance from generation to generation is known as the theory of the continuity of germ plasm.

Such appears to be, at least in part, the machinery of heredity. This understanding makes the germ substance perpetual and continuous, and explains why successive generations are alike. It does not explain, indeed, why an individual inherits from its parents, but why it is like its parents. While biologists are still in dispute over many problems connected with heredity, all are agreed to-day that this principle of the continuity of the heredity substance must be the basis of all attempts to understand the machinery of heredity. But plainly this whole process is a function of the cell machinery. While, therefore, the idea of the continuity of germ substance greatly simplifies our problem, we must acknowledge that once more we are thrown back upon the mysteries of the cell. Until we can more fully explain the cell machine we must recognize our inability to solve the fundamental question of why an individual is like its parents.

[Illustration: FIG. 50.—Diagram illustrating the principle of heredity.

A represents an egg of a starfish. From one half, the unshaded portion, develops the starfish of the next generation, B. The other is distributed without change in the ovaries, ov, of the individual, B. From these ovaries arises the next egg, A', with its germ plasm. This germ plasm is evidently identical with that in A, since it is merely a bit of the same handed down through the individual, B. In the development of the next generation the process is repeated, and hence B' will be like B, and the third generation of eggs identical with the first and second. The undifferentiated part of the germ plasm is thus simply handed on from one generation to the next.]

But plainly reproduction and heredity, as we have thus far considered them, will be unable to account for the slow modification of the machine; for in accordance with the facts thus far outlined, each generation would be precisely like the last, and there would be no chance for development and change from generation to generation. If the individual is simply the unfolding of the powers possessed by a bit of germ plasm, and if this germ plasm is simply handed on from generation to generation, the successive generations must of necessity be identical. But the living machine has been built by changes in the successive generation, and hence plainly some other factor is needed. This factor is variation.

Variation.—Variation is the principle that produces modification of type. Heredity, as just explained, would make all generations alike. But nothing is more certain than that they are not alike. The fact of variation is patent on every side, for no two individuals are alike. Successive generations differ from each other in one respect or another. Birds vary in the length of their bills or toes; butterflies, in their colours; dogs, in their size and shape and markings; and so on through an endless category. Plants and animals alike throughout nature show variations in the greatest profusion. It is these variations which must furnish us with the foundation of the changes which have gradually built up the living machine.

Of the fact of these variations there is no question, and the matter need not detain us. Every one has had too many experiences to ask for proof. Of the nature of the variations, however, there are some points to be considered which are very germane to our subject. In the first place, we must notice that these variations are of two kinds. There is one class which is born with the individual, so that they are present from the time of birth. In saying that these variations are born with the individual we do not necessarily mean that they are externally apparent at birth. A child may inherit from its parents characters which do not appear till adult life. For example, a child may inherit the colour of its father's hair, but this colour is not apparent at birth. It appears only in later life, but it is none the less an inborn character. In the same way, we may have many inborn variations among individuals which do not make themselves seen until adult life, but which are none the less innate. The offspring of the same parents may show decided differences, although they are put under similar conditions, and such differences are of course inherent in the nature of the individual. Such variations are called congenital variations.

There is, however, a second class of variations which are not born in the individual, but which arise as the result of some conditions affecting its after-life. The most extreme instances of this kind are mutilations. Some men have only one leg because the other has been lost by accident. Here is a variation acquired as the result of circumstances. A blacksmith differs from other members of his race in having exceptionally large arm muscles; but here, again, the large muscles have been produced by use. A European who has lived under a tropical sun has a darkened skin, but this skin has evidently been darkened by the action of the sun, and is quite a different thing from the dark skin of the dark races of men. In such instances we have variations produced in individuals as the result of outside influences acting upon them. They are not inborn, but are secondarily acquired by each individual. We call them acquired variations.

It is not always possible to distinguish between these two types of variation. Frequently a character will be found in regard to which it is impossible to determine whether it is congenital or acquired. If a child is born under the tropical sun, how can we tell whether its dark skin was the result of direct action of the sun on its own skin, or was an inheritance from its dark-skinned parents? We might suppose that this could be answered by taking a similar child, bringing it up away from the tropical sun, and seeing whether his skin remained dark. This would not suffice, however; for if such a child did then develop a white skin, we could not tell but that this lighter-coloured skin had been produced by the direct bleaching effect of the northern climate upon a skin which otherwise would have been dark. In other words, a conclusive answer can not here be given. It is not our purpose, however, to attempt to distinguish between these two kinds of variations, but simply to recognize that they occur.

Our next problem must be to search for an explanation of these variations. With the acquired variations we have no particular trouble, for they are easily explained as due to the direct action of the environment upon animals. One of the fundamental characters of the living protoplasm (using the word now in its widest sense) is its extreme instability. So unstable is it that any disturbing influence will affect it. If two similar unicellular organisms are placed under different conditions they become unlike, since their unstable protoplasm is directly affected by the surrounding conditions. With higher animals the process is naturally a little more complicated; but here, too, they are easily understood as part of the function of the machine. One of the adjustments of the machine is such that when any organ is used more than usual the whole machine reacts in such a way as to send more blood to this special organ. The result is a change in the nutrition of the organ and a corresponding variation in the individual. Thus acquired variations are simply functions of the action of the machine.

Congenital variations, however, can not receive such an explanation. Being born with the individual, they can not be produced by conditions affecting him, but rather to something affecting the germ plasm from which he sprung. The nature of the germ plasm controls the nature of the individual, and congenital variations must consequently be due to its variations. But it is not so easy to see how this germ plasm can undergo variation. The conditions which surround the individual would affect its body, but it is not easy to believe that they would affect the germinal substance. Indeed, it is not easy to see how any external conditions can have influence upon this germinal material if it is not an active part of the body, but is simply stored within it for future use in reproduction. How could any changes in the environment of the individual have any effect upon this dormant material stored within it? But if we are correct in regarding this germ material in the reproductive bodies as the basis of heredity and the guiding force in development, then it follows that the only way in which congenital variations can occur is by some variations in the germ plasm. If a child developed from germ plasm identical with that from which its parents developed, it would inherit identical characters; and if there are any congenital variations from its parents, they must be due to some variations in the germ plasm. In other words, in order to explain congenital variations we must account for variations in the germ plasm.

Now, there are two methods by which we may suppose that these variations in the germ may arise. The first is by the direct influence upon the germ plasm of certain unknown external conditions. The life substance of organisms is always very unstable, and, as we have seen, acquired variations are caused by external influences directly affecting it. Now, the hereditary material is also life substance, and it is plainly a possibility for us to imagine that this germ material is also subject to influences from the conditions surrounding it. That such variations do occur appears to be hardly doubtful, although we do not know what sort of influences can produce them. If the germ plasm is wholly stored within the reproductive gland, it is certainly in a position to be only slightly affected by surrounding conditions which affect the animal. We can readily understand that the use of an organ like the arm will affect it in such a way as to produce changes in its protoplasm, but we can hardly imagine that such use of the arm would produce any change in the hereditary substance which is stored in the reproductive organs. External conditions may thus readily affect the body, but not so readily the germ material. Even if such material is distributed more or less over the body instead of being confined to the reproductive glands, as some believe, the difficulty is hardly lessened. This difficulty of understanding how the germ plasm can be affected by external conditions has led one school of biologists to deny that it is subject to any variation by external conditions, and hence that all modification of the germ plasm must come from some other source. Probably no one, however, holds this position to-day, and it is the general belief that the germ plasm may be to some slight extent modified by external conditions. Of course, if such variations do occur in the germ plasm they will become congenital variations of the next generation, since the next generation is the unfolding of the germ plasm.

The second method by which the variations of germ plasm may arise is apparently of more importance. It is based upon the fact that, with all higher animals and plants at least, each individual has two parents instead of one. In our study of cells we have seen that the machinery of the cell is such that it requires in the ordinary process of reproduction the union of germinal material from two different individuals to produce a cell which can develop into a new individual. As we have seen, the egg gets rid of half its chromosomes in order to receive an equal number from a male parent; and thus the fertilized egg contains chromosomes, and hence hereditary material, from two different individuals. Now, this sexual reproduction occurs very widely in the organic world. Among some of the lowest forms of unicellular organisms it is not known, but in most others some form of such union is universal. Now, here is plainly an abundant opportunity for congenital variations; for it is seen that each individual does not come from germ material identical with that from which either parent came, but from some of this material mixed with a similar amount from a different parent. Now, the two parents are never exactly alike, and hence the germ plasm which each contributes to the offspring will not be exactly alike. The offspring will thus be the result of the unfolding of a bit of germ plasm which will be different from that from which either of its parents developed, and these differences will result in congenital variations. Sexual reproduction thus results in congenital variations; and if congenital variations are necessary for the evolution of the living machine—and we shall soon see reason for believing that they are—we find that sexual reproduction is a device adopted for bringing out such congenital variations.

Inheritance of Variations.—The reason why congenital variations are needed for the evolution of the living machine is clear enough. Evanescent variations can have no effect upon this machine, for they would disappear with the individual in which they appeared. In order that they should have any influence in the process of machine building they must be permanent ones; or, in other words, they must be inherited from generation to generation. Only as such variations are transmitted by heredity can they be added to the structure of the developing machine. Therefore we must ask whether the variations are inherited.

In regard to the congenital variations there can be no difficulty. The very fact that they are congenital shows us that they have been produced by variations in the germ plasm, and as such they must be transmitted, not only to the next generation, but to all following generations, until the germ plasm becomes again modified. This germ plasm is handed on from generation to generation with all its variations, and hence the variations will be added permanently to the machine. Congenital variations are thus a means for permanently modifying the organism, and by their agency must we in large measure believe that evolution through the ages has taken place.

With the acquired variations the matter stands quite differently. We can readily understand how influences surrounding an animal may affect its organs. The increase in the size of the muscles of the blacksmith's arm by use we understand readily enough. But with our understanding of the machinery of heredity we can not see how such an effect can extend to the next generation. It is only the organ directly affected that is modified by external conditions. Acquired variations will appear in the part of the body influenced by the changed conditions. But the germ plasm within the reproductive glands is not, so far as we can see, subject to the influence of an increased use, for example, in the arm muscles. The germ material is derived from the parents, and, if it is simply stored in the individual, how could an acquired variation affect it? If an individual lose a limb his offspring will not be without a corresponding limb, for the hereditary material is in the reproductive organs, and it is impossible to believe that the loss of the limb can remove from the hereditary material in the reproductive glands just that part of the germ plasm which was designed for the production of the limb. So, too, if the germ plasm is simply stored in the individual, it is impossible to conceive any way that it can be affected by the conditions around the individual in such a way as to explain the inheritance of acquired variations. If acquired variations do not affect the germ plasm they cannot be inherited, and if the germ plasm is only a bit of protoplasmic substance handed down from generation to generation, we can not believe that acquired variations can influence it.

From such considerations as these have arisen two quite different views among biologists; and, while it is not our purpose to deal with disputed points, these views are so essential to our subject that they must be briefly referred to. One class of biologists adhere closely to the view already outlined, and insist for this reason that acquired variations can not under any conditions be inherited. They insist that all inherited variations are congenital, and due therefore to direct variations in the germ plasm, and that all instances of seeming inheritance of acquired variations are capable of other explanation. The other school is equally insistent that there are abundant instances of the inheritance of acquired characters, claiming that these proofs are so strong as to demand their acceptance. Hence this class of biologists insist that the explanation of heredity given as a simple handing down from generation to generation of a germ plasm is not complete, and that while it is doubtless the foundation of heredity, it must be modified in some way so as to admit of the inheritance of acquired characters. There is no question that has excited such a wide interest in the biological world during the last fifteen years as this one of the inheritance of acquired characters. Until about 1884 the question was not seriously raised. Heredity was known to be a fact, and it was believed that while congenital characters are more commonly inherited, acquired characters may also frequently be handed down from generation to generation. The facts which we have noted of the continuity of germ plasm have during the last fifteen years led many biologists to deny the possibility of the latter. The debate which arose has continued vigorously, and can not be regarded as settled at the present time. One result of this debate is clear. It has been shown beyond question that while the inheritance of congenital characters is the rule, the inheritance of acquired characters is at all events unusual. At the present time many naturalists would be inclined to think that the balance of evidence indicates that under certain conditions certain kinds of acquired characters may be inherited, although this is still disputed by others. Into this discussion we cannot enter here. The reason for referring to it at all is, however, evident. We are searching for nature's method of building machines. It is perfectly clear that variations among animals and plants are the foundations of the successive steps in advance made in this machine building, but of course only such variations as can be transmitted to posterity can serve any purpose in this development. If therefore it should prove that acquired characters can not be inherited, then we should no longer be able to look upon the direct influence of the surroundings as a factor in the machine building. We should then have nothing left except the congenital variations produced by sexual union, or the direct variation of the germ plasm as a factor for advance. If, however, it shall prove that acquired characters may even occasionally be inherited, then the direct effect of the environment upon the individual will serve as a decided assistance in our problem.

Here, then, we have before us the factors which have been concerned in the building of the living machine under nature's hands. Reproduction keeps in existence a constantly active, unstable, readily modified organism as a basis upon which to build. Variation offers constantly new modifications of the type, while heredity insures that the modifications produced in the machine by the influences which give rise to the variations shall be permanently fixed.

Method of Machine Building.—Natural Selection. The method by which these factors have worked together to build up the living machines is easily understood in its general aspects, although there are many details as yet unsolved. The general facts connected with the evolution of animals are matters of common knowledge. We need do no more than outline the subject, since it is well understood by all. The basis of the method is natural selection, which acts in this machine building something as follows:

The law of reproduction, as we have seen, produces new individuals with extraordinary rapidity, and as a result more individuals are born than can possibly find sustenance in the world. Hence only a few of the offspring of any animal or plant can live long enough to produce offspring in turn. The many must die that the few may live; and there is, therefore, a constant struggle among the individuals that are born for food or for room in the world. In this struggle for existence of course the weakest will go to the wall, while those that are best adapted for their place in life will be the ones to get food, live, and reproduce their kind. This is at all events true among the lower animals, although with mankind the law hardly applies. Now, among the individuals that are born there will be no two exactly alike, since variations are universal, many of which are congenital and thus born with the individual and transmitted by inheritance. Clearly enough those animals that have a variation which makes them a little better adapted for the struggle will be the ones to live and hence to produce offspring, while those without such advantage will be the ones to die. We may suppose, for example, that some of the individuals had longer necks than the average. In time of scarcity of food these individuals would be able to get food that the short-necked individuals could not reach. Hence in times of famine the long-necked individuals would be the ones to survive. Now if this peculiarity were a congenital variation it would be already represented in the germ plasm, and consequently it would be inherited by the next generation. The short-necked individuals being largely destroyed in this struggle for food, it would follow that the next generation would be a little better off than the last, since all would inherit this tendency toward a long neck. A few generations would then see the disappearance of all individuals which did not show either this or some other corresponding advantage, and in this way the lengthened neck would be added permanently as a part of the machine. When this time came this peculiarity would no longer give its possessors any advantage over its rivals, since all would possess it. Now, therefore, some new variation would in the same way determine which animals should live and which should die in the struggle, and in time a new modification would be added to the machine. And thus this process continues, one variation after another being added, until the machine is slowly built into a more and more complicated structure, always active but with a constantly increasing efficiency. The construction is a natural one. A mixing of germ plasm in sexual reproduction or some other agencies produce congenital variations; natural selection acting upon the numerous progeny selects the best of the new variations, and heredity preserves and hands them down to posterity.

All students of whatever school recognize the force of this principle and look upon natural selection as an efficient agency in machine building. It is probably the most fundamental of the external laws that have guided the process. There are, however, certain other laws which have played a more or less subordinate part. The chief of these are the influence of migration and isolation, and the direct influence of the environment. Each of these laws has its own school of advocates, and each has been given by its advocates the chief role in the process of machine building.

Migration and Isolation.—The production of the various types of machines has been undoubtedly facilitated by the migrations of animals and the isolation of different groups of descendants from each other by various natural barriers. The variations which occur in organisms are so great that they would sometimes run into abnormal structures were it not for the fact that sexual reproduction constantly tends to reduce them. In an open country where animals and plants interbreed freely, it will commonly happen that individuals with certain peculiarities will mate with others without such peculiarities, and the offspring will therefore inherit the peculiarity not in increased degree but in decreased degree. This constant interbreeding of individuals will tend to prevent the formation of many modifications in the machine which become started by variations. Now plainly if some such individuals, with a peculiar variation, should migrate into a new territory or become isolated from their relatives which do not have similar variations, these individuals will be obliged to breed with each other. The result will be that the next generation, arising thus from two parents each of which shows the same variation, will show it also in equal or increased degree. Migrations and isolations will thus tend to fix in the machine variations which sexual union or other influences inaugurate. Now in the history of the earth's surface there have been many changes which tend to bring about such migration and isolations, and this factor has doubtless played a more or less important part in the building of the machines. How great a part we cannot say, nor is it necessary for our purpose to decide; for in all these cases the machine building has only been the result of the hereditary transmission of congenital variation under certain peculiar conditions. The fundamental process is the same as already considered, only the details of its working being in question.

Direct Influence of the Environment.—Under this head we have a subject of great importance. It is an undoubted fact that the environment has a very decided effect upon the machine. These direct effects of the environment are very positive and in great variety. The tropical sun darkens the human skin; cold climate stunts the growth of plants; lack of food dwarfs all animals and plants, and hundreds of other similar examples could be selected. Another class of similar influences are those produced by use and disuse. Beyond question the use of an organ tends to increase its size, and disuse to decrease it. Combats of animals with each other tend to increase their strength, flight from enemies their running powers, etc.

Now all these effects are direct modifications of the machine, and if they are only transmitted to following generations so as to become permanent modifications, they will be most important agencies in the machine building. If, on the other hand, they are not transmitted by heredity, they can have no permanent effect. We have here thus again the problem of the inheritance of acquired characters. We have already noticed the uncertainty surrounding this subject, but the almost universal belief in the inheritance of such characters requires us to refer to it again. It is uncertain whether such direct effects have any influence upon the offspring, and therefore whether they have anything to do with this machine building. Still, there are many facts which point strongly in this direction. For example, as we study the history of the horse family we find that an originally five-toed animal began to walk more and more on its middle toe, in such a way that this toe received more and more use, while the outer toes were used less and less. Now that such a habit would produce an effect upon the toes in any generation is evident; but apparently this influence extended from generation to generation, for, as the history of the animals is followed, it is found that the outer toes became smaller and smaller with the lapse of ages, while the middle one became correspondingly larger, until there was finally produced the horse with its one toe only on each foot. Now here is a line of descent or machine building in the direct line of the effects of use and disuse, and it seems very natural to suppose that the modification has been produced by the direct effect of the use of the organs. There are many other similar instances where the line of machine building has been quite parallel to the effects of use and disuse. If, therefore, acquired characters can be inherited to any extent, we have, in the direct influences of the environment an important agency in machine building. This direct effect of the conditions is apparently so manifest that one school of biologists finds in it the chief cause of the variations which occur, telling us that the conditions surrounding the organism produce changes in it, and that these variations, being handed down to subsequent generations, constitute the basis of the development of the machine. If this factor is entirely excluded, we are driven back upon the natural selection of congenital variations as the only kind of variations which can permanently effect the modification of the machine.