The Doctrine of Evolution: Its Basis and Its Scope

by Henry Edward Crampton

IV

EVOLUTION AS A NATURAL PROCESS

The purpose of the discussions up to this point has been to present the reasons drawn from the principal classes of zoölogical facts for believing that living things have transformed naturally to become what they now are. Even if it were possible to make an exhaustive analysis of all of the known phenomena of animal structure, development, and fossil succession, the complete bodies of knowledge could not make the evolutionary explanation more real and evident than it is shown to be by the simple facts and principles selected to constitute the foregoing outline. We have dealt solely with the evidences as to the fact of evolution; and now, having assured ourselves that it is worth while to so do, we may turn to the intelligible and reasonable evidence found by science which proves that the familiar and everyday "forces" of nature are competent to bring about evolution if they have operated in the past as they do to-day. Investigation has brought to light many of the subsidiary elements of the whole process, and these are so real and obvious that they are simply taken for granted without a suspicion on our part of their power until science directs our attention to them.

For one reason or another, those who take up this subject for the first time find it difficult to banish from their minds the idea that evolution, even if it ever took place, has been ended. They think it futile to expect that a scrutiny of to-day's order can possibly find influences powerful enough to have any share in the marvelous process of past evolution demonstrated by science. The naturalists of a century ago held a similar opinion regarding the earth, viewing it as an immutable and unchanged product of supernatural creation, until Lyell led them to see that the world is a plastic mass slowly altering in countless ways. It is no more true that living things have ceased to evolve than that mountains and rivers and glaciers are fixed in their final forms; they may seem everlasting and permanent only because a human life is so brief in comparison with their full histories. Like the development of a continent as science describes it, the origin of a new species by evolution, its rise, culmination, and final extinction may demand thousands of years; so that an onlooker who is himself only a conscious atom of the turbulent stream of evolving organic life does not live long enough to observe more than a small fraction of the whole process. Therefore living species seem unchanged and unchangeable until a conviction that evolution is true, and a knowledge of the method of science by which this conviction is borne upon one, guide the student onwards in the further search for the efficient causes of the process.

The biologist employs the identical methods used by the geologist in working out the past history of the earth's crust. The latter observes the forces at work to-day, and compares the new layers of rock now being formed with the strata of deeper levels; these are so much alike that he is led to regard the constructive influences of the past as identical with those he can now watch at work. Similarly the biologist must first learn, as we have done, the principles of animal construction and development, and of other classes of zoölogical facts, and then he must turn his attention from the dead object of laboratory analysis to the workings of organic machines. The way an organism lives its life in dynamic relations to the varied conditions of existence, as well as the mutual physiological relations of the manifold parts of a single organism, reveal certain definite natural forces at work. Therefore his next task is to compare the results accomplished by these factors in the brief time they may be seen in operation with the products of the whole process of organic evolution, to learn, like the geologist in his sphere, that the present-day natural forces are able to do what reason says they have done in the past.

When the subject of inquiry was the reality of evolution, it was perhaps surprising to find that even the most familiar animals like cats and frogs provided adequate data for science to use in formulating its principles. So it is with the matter of method; it is unnecessary to go beyond the observations of a day or a week of human life to find forces at work, as real and vital as animal existence and organic life themselves. This is true, because evolution is true, and because the lives of all creatures follow one consistent law. Our task is therefore much more simple than most people suppose it to be; let us look about us and classify what we may observe, increasing our knowledge from the wide array of equally natural facts supplied by the biologist.

The analogies of the steamship and the locomotive proved useful at many times during the discussion of the fact of evolution, and even in the present connection they will still be of service. The evolution of these dead machines has been brought about by man, who, as an element of their environment, has been their creator as well as the director of their historical transformations. The result of their changes has been greater efficiency and better adjustment or adaptation to certain requirements fixed by man himself. The whole process of improvement has been one, in brief, of trial and error; new inventions have often been worthless, and they have been relegated to the scrap-heap, while the better part has been finally incorporated in the type machine. In brief, then, the important elements in the evolution of these examples have been three; first, adaptation, second, the origination of new parts, and third, the retention of the better invention.

Are the creatures of the living world so constituted that biological equivalents of these three essential elements of mechanical evolution can be found? Are organisms adapted to the circumstances controlling their lives, and are they capable of changing naturally from generation to generation, and of transmitting their qualities to their offspring? These are definite questions that bring us face to face with the fundamental problems relating to the dynamics or workings of evolution. We need not ask for or expect to find complete answers, for we know that it is impossible to obtain them. But we may expect to accomplish our immediate object, which is to see that evolution is natural. Our attention must be concentrated upon the three biological subjects of adaptation, variation, and inheritance, and we must learn why science describes them as real organic phenomena and the results of natural causes.

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At the very outset, when the general characteristics of living things were considered, much was said on the subject of adaptation as a universal phenomenon of nature. It was not contended that perfection is attained by any living mechanism, but it was held that no place exists in nature for an organism that is incapable of adjusting itself to the manifold conditions of life. A modus vivendi must be established and some satisfactory degree of adaptation must be attained, or else an animal or a species must perish. With this fundamental point as a basis, we look to nature for two kinds of natural processes or factors, first, those which may originate variations as primary factors,—the counterparts of human ingenuity and invention in the case of locomotive evolution,—and the secondary factors of a preservative nature which will perpetuate the more adaptive organic changes produced by the first influences; it is clear that the latter are no less essential for evolution than the first causes for the appearance of variations.

The term "variation" is employed for the natural phenomenon of being or becoming different. It is an obvious fact that no child is ever exactly like either of its parents or like any one of its earlier ancestors; while furthermore in no case does an individual resemble perfectly another of its own generation or family. This departure from the parental condition, and the lack of agreement with others even of its closest blood-relatives, are two familiar forms of variation. As a rule, the degree to which a given organism is said to vary in a given character is most conveniently measured by the difference between its actual condition and the general average of its species, even though there is no such thing as a specimen of average nature in all of its qualities. In brief, then, variation means the existence of some differences between an individual and its parents, its fraternity, and, in a wider sense, all others of its species.

Passing now to the causes of variation, all of the countless deviations of living things can be referred to three kinds of primary factors; namely, the environmental, functional, and congenital influences that work upon the organism in different ways and at different times during its life. We shall learn that the evolutionary values of these three classes are by no means equal, but we take a long step forward when we realize that among the things we see every day are facts demonstrating the reality of three kinds of natural powers quite able to change the characters of organic mechanisms.

The "environment" of an organism is everything outside the creature itself. In the case of an animal it therefore includes other members of its own kind, and other organisms which prey upon its species or which serve it as food, as well as the whole series of inorganic influences which first come to mind when the term is used. For example, the environment of a lion includes other lions which are either members of its own family, or else, if they live in the same region, they are its more or less active rivals and competitors. In the next place, other kinds of animals exist whose lives are intimately related to the lion's life, such as the antelopes or zebras that are preyed upon, and the human hunter to whom the lion itself may fall a victim. In addition, there are the contrasted influences of inorganic nature which demand certain adjustments of the lion's activities. Light and darkness, heat and cold, and other factors have their direct and larger or smaller effects upon the life of a lion, although these effects are less obvious in this instance than in the case of lower organisms.

The reality of variations due to the inorganic elements of the environment is everywhere evident. Those who have spent much time in the sun are aware that sunburn may result as a product of a factor of this class. The amount of sunlight falling upon a forest will filter through the tree-tops so as to cause some of the plants beneath to grow better than others, thus bringing about variations among individuals that may have sprung from the myriad seeds of a single parent plant. In times of prolonged drought, plants cannot grow at the rate which is usual and normal for their species, and so many variations in the way of inhibited development may arise.

Then there are the variations of a second class, more complex in nature than the direct effects of environment,—namely, the functional results of use and disuse. A blacksmith uses his arm muscles more constantly than do most other men, and his prolonged exercise leads to an increase of his muscular capacity. All of the several organic systems are capable of considerable development by judicious exercise, as every one knows. If the functional modifications through use were unreal, then the routine of the gymnasium and the schoolroom would leave the body and the mind as they were before. Furthermore, we are all familiar with the opposite effects of disuse. Paralysis of an arm results in the cessation of its growth. When a fall has injured the muscles and nerves of a child's limb, that structure may fail to keep pace with the growth of the other parts of the body as a result of its disuse. These are simple examples of a wide range of phenomena exhibited everywhere by animals and even by the human organism, demonstrating the plasticity of the organic mechanism and its modification by functional primary factors of variation.

But by far the greater number of variations seem to be due to the so-called congenital causes, which are sharply contrasted with the influences of the first and second classes. It is quite true that the influences of the third class cannot be surely and directly demonstrated like the others, but however remote and vague they themselves may appear to be, their effects are obvious and real, while at the same time their effects are to be clearly distinguished from the products of the other two kinds. Congenital factors reside in the physical heritage of an organism, and their results are often evident before an individual is subjected to environmental influences and before it begins to use its various organs. For example, it is a matter of common observation that a child with light hair and blue eyes may have dark-eyed and brown-haired parents. The fact of difference is a phenomenon of variation; the causes for this fact cannot be found in any other category than that comprising the hereditary and congenital influences of parent upon offspring. How the effect is produced by such causes is less important in the present connection than the natural fact of congenital variation. Science, however, has learned much about the causes in question, as we shall see at a later point.

Thus the first step which is necessary for an evolution and transformation of organic mechanisms proves to be entirely natural when we give only passing attention to certain obvious phenomena of life. The fact of "becoming different" cannot be questioned without indicting our powers of observation, and we must believe in it on account of its reality, even though the ultimate analysis of the way variations of different kinds are produced remains for the future.

Having learned that animals are able to change in various ways, the next question is whether variations can be transmitted to future generations through the operation of secondary factors. Long ago Buffon held that the direct effects of the environment are immediately heritable, although the mode of this inheritance was not described; it was simply assumed and taken for granted. Thus the darker color of the skin of tropical human races would be viewed by Buffon as the cumulative result of the sun's direct effects. Lamarck laid greater stress upon the indirect or functional variations due to the factors of use and disuse, and he also assumed as self-evident that such effects were transmissible as "acquired characters." This expression has a technical significance, for it refers to variations that are added during individual life to the whole group of hereditary qualities that make any animal a particular kind of organism. If evolution takes place at all, any new kind of organism originating from a different parental type must truly acquire its new characteristics, but few indeed of the variations appearing during the lifetime of an animal owe their origin to the functional and environmental influences, whose effects only deserve the name of "acquired characters" in the special biological sense.

In sharp contrast to Lamarckianism, so called,—although it did not originate in the mind of the noted man of science whose name it bears,—is the doctrine of natural selection, first proposed in its full form by Charles Darwin. This doctrine presents a wholly natural description of the method by which organisms evolve, putting all of the emphasis upon the congenital causes of variation, although the reality of other kinds of change is not questioned. But the contrast between Darwinism and the other descriptions of secondary factors can best be made after a somewhat detailed discussion of the former, which has gained the adherence of the majority of the naturalists of to-day. However, we must not pass on without pointing out that however much the explanations given by various men of science may differ, they all agree in expressly recognizing the complete naturalness of the secondary as well as of the primary factors of evolution.

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The doctrine of natural selection forms the best basis for the detailed discussion of the way evolution has come about in the past and how it is going on to-day. This is true because it was the first description of nature's program to carry conviction to the scientific world, and because its major elements have stood the test of time as no other doctrine has done. Much has been added to our knowledge of natural processes during post-Darwinian times, and new discoveries have supplemented and strengthened the original doctrine in numerous ways, although they have corrected certain of the minor details on the basis of fuller investigation.

At the outset it must be clearly understood that Darwin's doctrine is concerned primarily with the method and not with the evidences as to the actual fact of evolution. Most of those who are not familiar with the principles of science believe that Darwin discovered this process; but their opinion is not correct. The reality of natural change as a universal attribute of living things had been clearly demonstrated long before Darwin wrote the remarkable series of books whose influence has been felt outside the domains of biology and to the very confines of organized knowledge everywhere. The "Origin of Species" was published in 1859, and only the last of its fourteen chapters is devoted to a statement of the evidence that evolution is true. In this volume Darwin presented the results of more than twenty-five years of patient study of the phenomena of nature, utilizing the observations of wild life in many regions visited by him when he was the naturalist of the "Beagle" during its famous voyage around the world. He also considered at length the results of the breeder's work with domesticated animals, and he showed for the first time that the latter have an evolutionary significance. Because his logical assembly of wide series of facts in this and later volumes did so much to convince the intellectual world of the reasonableness of evolution, Darwin is usually and wrongly hailed as the founder of the doctrine. It is interesting to note in passing that Alfred Russel Wallace presented a precisely similar outline of nature's workings at about the same time as the statement by Darwin of his theory of natural selection. But Wallace himself has said that the greater credit belongs to the latter investigator who had worked out a more complete analysis on the basis of far more extensive observation and research.

The fundamental point from which the doctrine of natural selection proceeds is the fact that all creatures are more or less perfectly adapted to the circumstances which they must meet in carrying on their lives; this is the reason why so much has been said in earlier connections regarding the universal occurrence of organic adaptation. An animal is not an independent thing; its life is intertwined with the lives of countless other creatures, and its very living substance has been built up out of materials which with their endowments of energy have been wrested from the environment. Every animal, therefore is engaged in an unceasing struggle to gain fresh food and new energy, while at the same time it is involved in a many-sided conflict with hordes of lesser and greater foes. It must prevail over all of them, or it must surrender unconditionally and die. There is no compromise, for the vast totality we individualize as the environment is stern and unyielding, and it never relents for even a moment's truce.

To live, then, is to be adapted for successful warfare; and the question as to the mode of origin of species may be restated as an inquiry into the origin of the manifold adaptations by which species are enabled to meet the conditions of life. Why is adaptation a universal phenomenon of organic nature?

The answer to this query given by Darwinism may be stated so simply as to seem almost an absurdity. It is, that if there ever were any unadapted organisms, they have disappeared, leaving the world to their more efficient kin. Natural selection proves to be a continuous process of trial and error on a gigantic scale, for all of living nature is involved. Its elements are clear and real; indeed, they are so obvious when our attention is called to them that we wonder why their effects were not understood ages ago. These elements are (1) the universal occurrence of variation, (2) an excessive natural rate of multiplication, (3) the struggle for existence entailed by the foregoing, (4) the consequent elimination of the unfit and the survival of only those that are satisfactorily adapted, and (5) the inheritance of the congenital variations that make for success in the struggle for existence. It is true that these elements are by no means the ultimate causes of evolution, but their complexity does not lessen their validity and efficiency as the immediate factors of the process.

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Taking up the first proposition, we return to the subject of variation that has been discussed previously for the purpose of demonstrating its reality. The observations of every day are enough to convince us that no two living things are ever exactly alike in all respects. The reason is that the many details of organic structure are themselves variable, so that an entire organism cannot be similar to another either in material or in functional regards, while furthermore it would be impossible for an animal to be related to environmental circumstances in the same way as another member of its species unless it was possible for two things to occupy the same space at the same time! Individual differences in physical constitution are displayed by any litter of kittens, with identical parents; it needs only a careful examination to find the variations in the shape of the heads, the length of their tails, and in every other character. Sometimes the differences are less evident in physical qualities than in disposition and mental make-up, for such variations can be found among related kittens just as surely as among the children belonging to a single human family.

Not only do all organisms vary, but they seem to vary in somewhat similar ways. While modern investigations have thrown much light upon the relations between variations and their causes, of particular value in the case of the congenital phenomena, the greatest advance since Darwin's time consists in the demonstration by the naturalists who have employed the laborious methods of statistical analysis that the laws according to which differences occur are the same where-ever the facts have been examined. A single illustration will suffice to indicate the general nature of this result. If the men of a large assemblage should group themselves according to their different heights in inches, we would find that perhaps one half of them would agree in being between five feet eight inches and five feet nine inches tall. The next largest groups would be those just below and above this average class,—namely, the classes of five feet seven to eight inches and five feet nine to ten inches. Fewer individuals would be in the groups of five feet five to six inches and five feet ten to eleven inches, and still smaller numbers would constitute the more extreme groups on opposite sides of these. If the whole assemblage comprised a sufficient number of men, it would be found that a class with a given deviation from the average in one direction would contain about the same number of individuals as the class at the same distance from the average in the opposite direction. Taking into account the relative numbers in the several classes and the various degrees to which they depart from the average, the mathematician describes the whole phenomenon of variation in human stature by a concise formula which outlines the so-called "curve of error." From his study of a thousand men, he can tell how many there would be in the various classes if he had the measurements of ten thousand individuals, and how many there would be in the still more extreme classes of very short and very tall men which might not be represented among one thousand people.

It is not possible to explain why variation should follow this or any other mathematical law without entering into an unduly extensive discussion of the laws of error. The mathematicians themselves tell us in general terms that the observations they describe so simply by their formulæ follow as the result of so-called chance, by which they mean that the combined operation of numerous, diverse, and uncorrelated factors brings about this result, and not, of course, that there is such a thing as an uncaused event or phenomenon.

Whenever any extensive series of like organisms has been studied with reference to the variations of a particular character, the variations group themselves so as to be described by identical or similar curves of error. It is certainly significant that this is true for such diverse characters, cited at random from the lists of the literature, as the number of ray-flowers of white daisies, the number of ribs of beech leaves, and of the bands upon the capsules of poppies, for the shades of color of human eyes, for the number of spines on the backs of shrimps, and for the number of days that caterpillars feed before they turn into pupæ.

To summarize the foregoing facts, we have learned that variation is universal throughout the living world, and that the primary factors causing organic difference—the counterparts of human ingenuity in the case of dead mechanisms—are the natural influences of the environment, of organic physiological activity, and of congenital inheritance. These factors are accorded different values in the evolution of new species, as we may see more clearly at a later juncture, but the essential point here is that they are not unreal, although they may not as yet be described by science in final analytical terms.

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We come now to the second element of the whole process of evolution, namely, what we may call overproduction or excessive multiplication. Like variation and so many other phenomena of nature, this is so real and natural that it escapes our attention until science places it before us in a new light. The normal rate of reproduction in all species of animals is such that if it were unchecked, any kind of organism would cumber the earth or fill the sea in a relatively short time. That this is universally true is apparent from any illustration that might be selected. Let us take the case of a plant that lives for a single year, and that produces two seeds before it withers and dies; let us suppose that each of these seeds produces an adult plant which in its turn lives one year and forms two seeds. If this process should continue without any interference, the twentieth generation after as many years would consist of more than one million descendants of the original two-seeded annual plant, provided only that each individual of the intervening years should live a normal life and should multiply at the natural rate. But such a result as this is rendered impossible by the very nature which makes annual plants multiply in the way they do. Let us take the case of a pair of birds which produce four young in each of four seasons. Few would be prepared for the figures enumerating the offspring of a single pair of birds at the end of fifteen years, if again all individuals lived complete and normal lives: at the end of the time specified there would be more than two thousand millions of descendants. The English sparrow has been on this continent little more than fifty years; it has found the conditions in this country favorable because few natural enemies like those of its original home have been met, and as a consequence it has multiplied at an astounding rate so as to invade nearly all parts of North America, driving out many species of song birds before it. About twenty years ago David Starr Jordan wrote that if the English sparrow continued to multiply at the natural rate of that time, in twenty years more there would be one sparrow to every square inch of the state of Indiana; but of course nature has seen to it that this result has not come about. A single conger-eel may produce fifteen million eggs in a single season, and if this natural rate of increase were unchecked, the ocean would be filled solid with conger-eels in a few years. Sometimes a single tapeworm, parasitic in the human body, will produce three hundred million embryos; the fact that this animal is relatively rare diverts our attention from the alarming fertility of the species and the excessive rate of its natural increase. Perhaps the most amazing figures are those established by the students of bacteria and other micro-organisms. Many kinds of these primitive creatures are known where the descendants of a single individual will number sixteen to seventeen millions after twenty-four hours of development under ordinarily favorable conditions. Though a single rodlike individual taken as a starting-point may be less than one five-thousandth of an inch in length, under natural circumstances it multiplies at a rate which within five days would cause its descendants to fill all the oceans to the depth of one mile. This is a fact, not a conjecture; the size of one organism is known, and the rate of its natural increase is known, so that it is merely a matter of simple arithmetic to find out what the result would be in a given time.

Even in the case of those animals that reproduce more slowly, an overcrowding of the earth would follow in a very short time. Darwin wrote that even the slow-breeding human species had doubled in the preceding quarter century. An elephant normally lives to the age of one hundred years; it begins to breed at the age of thirty, and usually produces six young by the time it is ninety. Beginning with a single pair of elephants and assuming that each individual born should live a complete life, only eight hundred years would be requisite to produce nineteen million elephants; a century or two more and there would be no standing room for the latest generation of elephants. It is only too obvious that such a result is not realized in nature, but it is on account of other natural checks, and not because the natural rate of reproductive increase is anything but excessive.

The third element of the process of natural selection is the struggle for existence which is to a large extent the direct consequence of over-multiplication. Because nature brings more individuals into existence than it can support, every animal is involved in many-sided battles with countless foes, and the victory is sometimes with one and sometimes with another participant in the conflict. A survivor turns from one vanquished enemy only to find itself engaged in mortal combat with other attacking forces. Wherever we look, we find evidence of an unceasing struggle for life, and an apparently peaceful meadow or pond is often the scene of fierce battles and tragic death that escape our notice only because the contending armies are dumb.

A community of ants, often comprising more individuals than an entire European state, depends for its national existence upon its ability to prevail over other communities with which it may engage in sanguinary wars where the losses of a single battle may exceed those of Gettysburg. The developing conger-eels find a host of enemies which greatly deplete their numbers before they can grow even into infancy. An annual plant does not produce a million living offspring in twenty years because seeds do not always fall upon favorable soil, nor do they always receive the proper amount of sunlight and moisture, or escape the eye of birds and other seed-eating animals. These three illustrations bring out the fact that there are three classes of natural conditions which must be met by every living creature if it is to succeed in life. In detail, the struggle for existence is intra-specific, involving some form of competition or rivalry among the members of a single species; it is inter-specific, as a conflict is waged by every species with other kinds of living things; and finally it involves an adjustment of life to inorganic environmental influences. While it may seem unjustifiable to speak of heat and cold and sunlight as enemies, the direct effects produced by these forces are to be reckoned with no less certainty than the attacks of living foes.

The three divisions of the struggle for existence are so important not only in purely scientific respects, but also in connection with the analysis of human biology, that we may look a little further into their details, taking them up in the reverse order. Regarding the environmental influences, the way that unfavorable surroundings decimate the numbers of the plants of any one generation has already been noted, and it is typical of the vital situation everywhere. English sparrows are killed by prolonged cold and snow as surely as by the hawk. The pond in which bacteria and protozoa are living may dry up, and these organisms may be killed by the billion. Even the human species cannot be regarded as exempt from the necessity of carrying on this kind of natural strife, for scores and hundreds die every year from freezing and sunstroke and the thirsts of the desert. Unknown thousands perish at sea from storm and shipwreck, while the recorded casualties from earthquakes and volcanic eruptions and tidal waves have numbered nearly one hundred and fifty thousand in the past twenty-eight years. The effects of inorganic influences upon all forms of organic life must not be underestimated in view of such facts as these.

In the second place, the vital struggle includes the battles of every species with other kinds of living things whose interests are in opposition. The relations of protozoa and bacteria, conger-eels and other fish, English sparrows and hawks, plants and herbivorous animals, are typical examples of the universal conflict in which all organisms are involved in some way. Again it is only too evident that human beings must participate every day in some form of warfare with other species. In order that food may be provided for mankind the lives of countless wild organisms must be sacrificed in addition to the great numbers of domesticated animals reared by man only that they may be destroyed. The wolf and the wildcat and the panther have disappeared from many of our Eastern states where they formerly lived, while no longer do vast herds of bison and wild horses roam the Western prairies. Because one or another human interest was incompatible with the welfare of these animals they have been driven out by the stronger invaders.

That the victory does not always fall to the human contestant is tragically demonstrated by the effects of the incessant assaults upon man made by just one kind of living enemy,—the bacillus of tuberculosis. Every year more than one hundred and twenty-five thousand people of the United States die because they are unable to withstand its persistent attacks; five million Americans now living are doomed to death at the hands of these executioners, and the figures must be more than doubled to cover the casualties on the human side in the battles with the regiments of all the species of bacteria causing disease.

The competition between and among the individuals of one and the same species is the third part of the struggle for existence, and it is often unsurpassed in its ferocity. When two lion cubs of the same litter begin to shift for themselves, they must naturally compete in the same territory, and their contest is keener than that which involves either of them and a young lion born ten or fifteen miles away. The seeds of one parent plant falling in a restricted area will be engaged in a competitive struggle for existence that is much more intense than many other parts of nature's warfare. In brief, the intensity of the competition will be directly proportional to the similarity of two organisms in constitution and situation, and to the consequent similarity of vital welfare. The interests of the white man and the Indian ran counter to each other a few hundred years ago, and the more powerful colonists won. The assumption of the white man's burden too often demonstrates the natural effect of diversity of interest, and the domination of the stronger over the weaker. In any civilized community the manufacturer, farmer, financier, lawyer, and doctor must struggle to maintain themselves under the conditions of their total inorganic and social environments; and in so far as the object of each is to make a living for himself, they are competitors. But the contest becomes more absorbing when it involves broker and broker, lawyer and lawyer, financier and magnate, because in each case the contestants are striving for an identical need of success.

Although the severity of the conflict imposed by nature is somewhat modified in the case of social organisms, where community competes with community and nation with nation, no form of social organization has yet been developed where the individual contest carried on by the members of one community has been done away with. It is an inexorable law of nature that all living things must fight daily and hourly for their very lives, because so many are brought into the world with each new generation that there is not sufficient room for all. No organism can escape the struggle for existence except by an unconditional surrender that results in death. Everywhere we turn to examine the happenings of organic life we can find nothing but a wearisome warfare in which it is the ultimate and cruel lot of every contestant to admit defeat.

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What now are the results of variation, over-multiplication, and competition? Since some must die because nature cannot support all that she produces, since only a small proportion of those that enter upon life can find a foothold or successfully meet the hordes of their enemies, which will be the ones to survive? Surely those that have even the slightest advantage over their fellows will live when their companions perish. It is impossible that the result could be otherwise; it must follow inevitably from what has been described before. The whole process has its positive and its negative aspects: the survival of the fittest and the elimination of the unfit. Perhaps it would be more correct to say the more real element is the negative one, for those which are least capable of meeting their living foes and the decimating conditions of inorganic nature are the first to die, while the others will be able to prolong the struggle for a longer or shorter period before they too succumb. Thus the destruction of the unfit leaves the field to the better adapted, that is, to those that vary in such a way as to be completely or at least partially adapted to carry on an efficient life. In this way Darwinism explains the universal condition of organic adjustment, showing that it exists because there is no place in nature for the incompetent.

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Finally we come to the process of inheritance as viewed by Darwin, and its part in the production and perfection of new species. In every case, Darwin said, the efficiency or inefficiency of an animal depends upon its characteristics of an inherited or congenital nature. Variations in these qualities provide the array of more or less different individuals from which impersonal nature selects the better by throwing out first the inferior ones. An organism can certainly change in direct response to environmental influence or by the indirect results of use and disuse, but not unless it is so constituted by heredity as to be able to change adaptively. Therefore the final basis of success in life must be sought in the inherited constitutions of organic forms.

For the reason that the qualities which preserve an animal's existence are already congenital, they are already transmissible, as Darwin contended. Since his time much has been learned about the course of inheritance and its physical basis, and the new discoveries have confirmed the essential truth of Darwin's statement that the congenital characters only possess a real power in the evolution of species.

We must devote some time to the subject of inheritance at a later juncture, but before leaving the matter an additional point must be established here; the selective process deals immediately with congenital results, as the heritable characters that make for success or failure in life, but by doing this it really selects the group of congenital factors behind and antecedent to their effects. For example, an ape that survives because of its superior cunning, does so because it varies congenitally in an improved direction; and the factors that have made it superior are indirectly but no less certainly preserved through the survival of their results in the way of efficiency. Hereditary strains are thus the ultimate things selected through the organic constitutions that they determine and produce.

Natural selection, as the whole of this intricate process, is simply trial and error on a gigantic scale. Nature is such that thousands of varying individuals are produced in order that a mere handful or only one survivor may be chosen to bear the burden of carrying on the species for another generation. The effect of nature's process is judicial, as it were. We may liken the many and varied conditions of life to as many jurymen, before which every living thing must appear for judgment as to its fitness or lack of it. A unanimous verdict of complete or partial approval must be rendered, or an animal dies, for the failure to meet a single vital condition results in sure destruction. Of course, we cannot regard selection as involving anything like a primitive conscious choice. It is because we individualize all of the complex totality of the world as "Nature" with a capital N that so many people unconsciously come to think of it as a human-like personality. He who would go further and hold that all of nature is actually conscious and the dwelling-place of the supernatural ultimate, must beware of the logical results of such a view. What must we think of the ethical status of such a conscious power who causes countless millions of creatures to come into the world and ruthlessly compels them to battle with one another until a cruel and tragic death ends their existence?

But that is a metaphysical matter, with which we need not concern ourselves in this discussion; the important point is that among the everyday happenings of life are processes that are quite competent to account for the condition of adaptation exhibited by various animal forms. These processes are real and natural, not imaginative or artificial, and so they will remain even though it will become clear that much is still to be learned about the causes of variation and the course of biological inheritance. Darwin was the first to contend that natural selection is but a part of nature's method of accomplishing evolution. As such it is content to recognize variations and does not concern itself with the origin of modifications; it accepts the obvious fact that congenital variations are inherited, although it leaves the question as to how they are inherited for further examination. Because the doctrine of natural selection does not profess to answer all the questions propounded by scientific inquisitiveness, it must not be supposed that it fails in its immediate purpose of giving a natural explanation of how evolution may be partly accounted for.

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Before proceeding to the post-Darwinian investigations that have done so much to amplify the account of natural evolution, let us consider the contrasted explanation given by Lamarck and his followers. As we have stated earlier, Lamarckianism is the name given to the doctrine that modifications other than those due to congenital factors may enter into the heritage of a species, and may add themselves to those already combined as the peculiar characteristics of a particular species. Let us take the giraffe and its long neck as a concrete example. The great length of this part is obviously an adaptive character, enabling the animal to browse upon the softer leafy shoots of shrubs and trees. The vertebral column of the neck comprises just the same number of bones that are present in the short-necked relatives of this form, so that we are justified in accepting as a fact the evolution of the giraffe's long neck by the lengthening of each one of originally shorter vertebræ. The Lamarckian explanation of this fact would be that the earliest forms in the ancestry of the giraffe as such stretched their necks as they fed, and that this peculiar function with its correlated structural modification became habitual. The slight increase brought about by any single individual would be inherited and transmitted to the giraffes of the next generation; in other words, an individually acquired character would be inherited. The young giraffes of this next generation would then begin, not where their parents did, but from an advanced condition. Thus, by continued stretching of the neck and by continued transmission of the elongated condition, the great length of this part of the body in the modern giraffe would be attained.

The explanation of natural selection would be quite different. The Darwinian would say that all the young giraffes of any one generation would vary with respect to the length of the neck. Those with longer necks would have a slight advantage over their fellows in the extended sphere of their grazing territory. Being better nourished than the others, they would be stronger and so they would be more able to escape from their flesh-eating foes, like the lion. For the reason that their variation would be congenital and therefore already transmissible, their offspring would vary about the advanced condition, and further selection of the longer necked individuals would lead to the modern result.

The Lamarckian explanation encounters one grave difficulty which is not met by the second one, in so far as it demands some method by which a bodily change may be introduced into the stream of inheritance. So far, this difficulty has not been overcome, and the present verdict of science is that the transmission of characters acquired as the result of other than congenital factors is not proved. It would be unscientific to say that it cannot be proved in the future, but there are good a priori grounds for disbelief in the principle, while furthermore the results of experiments that have been undertaken to test its truth have been entirely negative. Rats and mice have had their tails cut off to see if this mutilation would have its effect upon their young, and though this has been done for more than one hundred successive generations the length of the tail has not been altered. Quite unconscious of the scientific problem, many human races have performed precisely similar experiments through centuries of time. In some classes of Chinese, the feet of young girls have been bound in such a way as to produce a small, malformed foot, but this has not resulted in any hereditary diminution in the size of the feet of Chinese females. Many other similar mutilations have been practised, as for example, the flattening of the skull of some North American Indians, but the deformity must be produced again with each recurring generation. One after another, the cases that were supposed to give positive evidence have been reinvestigated, with the result that has been stated above. It would seem, therefore, that heredity and congenital modification must play by far the greater part in the evolution of species.

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The doctrine of natural selection took form in the mind of Darwin mainly on account of three potent influences; these were, first, the geological doctrine of uniformitarianism proposed by Lyell, second, his own observations of wild life in many lands and his analysis of the breeder's results with domesticated animals, and third, the writings of Malthus dealing with overpopulation. As Darwin had read the works of Buffon, Lamarck, and Erasmus Darwin, his grandfather, who had written a famous treatise under the title of "Zoonomia," he was familiar with the evidences known in his student days tending to prove that organic evolution was a real natural process. Lyell's doctrine of uniform geological history made an early and deep impression upon his mind, and it led him to ask himself whether the efficient causes of past evolution might not be revealed by an analysis of the present workings of nature. As naturalist of the "Beagle" during its four years' cruise around the world, Darwin saw many new lands and observed varied circumstances under which the organisms of the tropics and other regions lived their lives. The fierce struggle for existence waged by the denizens of the jungle recalled to him the views of Malthus regarding overpopulation and its results. These and other influences led him to begin the remarkable series of note-books, from which it is interesting indeed to learn how the doctrine of natural selection began to assume a definite and permanent form in his mind, as year followed year, and evidence was added to evidence. And it is a valuable lesson to the student of science that for twenty-five years Darwin devoted all his time to the acquisition of facts before he gave his doctrine to the world in the famous "Origin of Species."

Darwin was particularly impressed by the way mankind has dealt with the various species of domesticated animals, and he was the first naturalist to point out the correspondence between the breeder's method of "artificial selection," and the world-wide process of natural selection. As every one knows, the breeder of race horses finds that colts vary much in their speed; discarding the slower animals, he uses only the swifter for breeding purposes, and so he perfects one type of horse. With other objects in view, the heavy draught horse, the spirited hackney, and the agile polo pony have been severally bred by exactly the same method. Among cattle many kinds occur, again the products of an artificial or human selection; hornless breeds have been originated, as well as others with wide-spreading or sharply curved horns; the Holstein has been bred for an abundant supply of milk as an object, while Jerseys and Alderneys excel in the rich quality of their milk. Various kinds of domesticated sheep and rabbits and cats also owe their existence to the employment of the selfsame method, unconsciously copied by man from nature; for men have found variations arising naturally among their domesticated animals, and they have simply substituted their practical purposes or their fancy for nature's criterion of adaptive fitness, preserving those that they wish to perfect and eliminating those unfitted to their requirements or ideas.

In the case of many of these and other examples, wild forms still occur which seem to be like the ancestral stock from which the domesticated forms have been produced. All the varied forms of dogs—from mastiff to toy-terrier, and from greyhound to dachshund and bulldog—find their prototypes in wild carnivora like the wolf and jackal. In Asia and Malaysia the jungle fowl still lives, while its domesticated descendants have altered under human direction to become the diverse strains of the barnyard, and even the peculiar Japanese product with tail feathers sometimes as long as twenty feet. That far-reaching changes can be brought about in a relatively short time is proved by the history of the game cock, which has nearly doubled in height since 1850, while at the same time its slender legs, long spurs, and other qualities have been perfected for the cruel sport for which it has been bred. Again, the wild rock pigeon seems to be the ancestral form from which the fantail and pouter and carrier-pigeon with their diverse characters have taken their origin.

It is true that some biologists have urged certain technical objections to the employment of domesticated animals and their history as analogies to the processes and results in wild nature. To my mind, however, artificial selection is truly a part of the whole process of natural selection. Man is but one element of the environment of tame forms, and his fancy or need is therefore one of the varied series of external criteria that must be met if survival is to be the result; failing this, elimination follows as surely as under the conditions of an area uninhabited or uninfluenced by mankind. Congenital variation is real, selection is real and the heredity of the more fit modification is equally real. Surely Darwin was right in contending that the facts of this class amplify the conception of natural selection developed on the basis of an analysis of wild life.

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Knowing the elements of the selective process, it is possible to analyze and to understand many significant phenomena of nature, and to gain a clearer conception of the results of the struggle for existence, especially when the human factor is involved. Let us see how much is revealed when the foregoing results are employed in a further study of some of nature's vital situations.

As a consequence of the many-sided struggle for existence, the interrelations of a series of species will approach a condition of equilibrium in an area where the natural circumstances remain relatively undisturbed for a long time. For example, among the field-mice of one generation, just as many individuals will survive as will be able to find food and to escape hereditary foes such as cats and snakes and owls. The number of owls, in their turn, will be determined by the number of available mice and other food organisms, as well as by the severity of the adverse circumstances that cause elimination of the less fit among the fledglings brought into the world. The vital chain of connections is sometimes astonishingly long and intricate. One remarkable illustration is given by Fiske, as an elaboration of an example cited by Darwin. He points out that the fine quality of the traditional roast beef of England is directly determined by the number of elderly spinsters in that country. The chain of circumstances is as follows: the quality of the clover fields, furnishing the best food for cattle, depends largely upon the visits to the clover-blossoms by wild bees, that accomplish the fertilization of the flowers by carrying pollen upon their bodies from one plant to another. Field-mice devour the young in the nests of these bees, so if there are few field-mice there will be many bees, and consequently better grazing for the cattle. The number of field-mice will vary according to the abundance of cats, and so the number of these domestic animals will exert an influence upon the whole foregoing chain of forms. But, as Fiske points out, cats are the favorite companions of elderly spinsters; therefore, if there are many of the latter, there will be more cats, fewer field-mice, more bees, richer clover fields, and finer cattle! Each link is real and the whole chain is a characteristic example of the countless ways that the natural destinies of living things are interrelated and intertwined.

The reality of such organic interrelationships is revealed with wonderful clearness in the numerous instances where some disturbing factor has altered one or another element of the balanced system. The invasion of the new world by Europeans has directly led to the partial or complete extinction of the tribes of Indians to whom the land formerly belonged; they have disappeared almost entirely from our state of New York, together with the bear and wolf and many other species of animals that formerly existed here. Wild horses and bison have also vanished before the advances of civilization and the alteration of their homes. Sometimes the extermination of one pest has resulted in an increase in the number of another through human interference with nature's equilibrium. In some of our Western states, a bounty was offered for the scalps of wolves, so as to lessen the number of these predatory foes of sheep. But when the wolves were diminished in number, their wild food-animals, the prairie dogs, found their lot much bettered, and they have multiplied so rapidly that in some places they have become even more destructive than the wolves.

One of the most remarkable illustrations is that of the rabbits introduced into Australia. This island continent was cut off from the surrounding lands long before the higher mammals evolved in far distant regions, so that the balance of nature was worked out without reference to animals like the rabbit. When the first of these were introduced they found a territory without natural enemies where everything was favorable. They promptly multiplied so rapidly that within a few years their descendants were numerous enough to eat up practically every green thing they could reach. Two decades ago, the single province of Queensland was forced to expend $85,000,000 in a vain effort to put down the rabbit plague. The remarkable statement has been made that in some places nature has taken a hand in causing a new type of rabbit to evolve. Finding the situation desperate, some of the animals have begun to develop into tree-climbing creatures. The animals exist in such numbers that the available food upon the ground is insufficient for all, and so some elimination results. But the young rabbits with longer claws, varying in this way on account of congenital factors, have an advantage over their fellows because they can climb some of the trees and so obtain food inaccessible to the others. If the facts are correctly reported, and if the process of selection on the basis of longer claws and the climbing habit is continued, the original type of animal is splitting up into a form that will remain the same and live upon the ground, and another that will be to all intents and purposes a counterpart of our familiar squirrel. All the evidence goes to show that squirrels have evolved from terrestrial rodents; if the data relating to Australian rabbits are correct, nature is again producing a squirrel-like animal by evolution in a region where the former natural situation has been interfered with by man.

The laws of biological inheritance have received close and deep study by numerous investigators of Darwinian and post-Darwinian times, because from the first it was clearly recognized that a complete description of nature's method of accomplishing evolution must show how species maintain the same general characteristics from generation to generation, and also how new qualities may be fixed in heredity as species transform in the course of time. Before our modern era in biology, the fact of inheritance was accepted as self-sufficient; now much is known that supplements and extends the incomplete account given by natural selection of the way evolution takes place.

It is not possible in the present brief outline to describe all the results of recent investigations, but some of them are too important to be passed over. Perhaps the most interesting one is that the laws of heredity seem to be the same for man and other kinds of living creatures, as proved by Galton and Pearson and many others who have dealt with such characters as human stature, human eye color, and an extensive series of the peculiarities of lower animals and even of plants.

The researches dealing with the physical basis of inheritance and its location in the organism have yielded the most striking and brilliant results. Darwin himself realized that the doctrine of natural selection was incomplete, as it accepted at its face value the inheritance of congenital racial qualities without attempting to describe the way an egg or any other germ bears them, and he endeavored to round out his doctrine of selection by adding the theory of pangenesis. According to this, every cell of every tissue and organ of the body produces minute particles called gemmules, which partake of the characters of the cells that produce them. The gemmules were supposed to be transported throughout the entire body, and to congregate in the germ-cells, which in a sense would be minute editions of the body which bears them, and would then be capable of producing the same kind of a body. If true, this view would lead to the acceptance of Lamarck's or even Buffon's doctrine, for changes induced in any organ by other than congenital factors could be impressed upon the germ-cell, and would then be transported together with the original specific characters to future generations. Darwin was indeed a good Lamarckian.

But the researches of post-Darwinians, and especially those of the students of cellular phenomena, have demonstrated that such a view has no real basis in fact. Many naturalists, like Naegeli and Wiesner, were convinced that there was a specific substance concerned with hereditary qualities as in a larger way protoplasm is the physical basis of life. It remained for Weismann to identify this theoretical substance with a specific part of the cell, namely, the deeply staining substance, or chromatin, contained in the nucleus of every cell. Bringing together the accumulating observations of the numerous cytologists of his time, and utilizing them for the development of his somewhat speculative theories, Weismann published in 1882 a volume called "The Germ Plasm," which is an immortal foundation for all later work on inheritance. The essential principles of the germ-plasm theory are somewhat as follows. The chromatin of the nucleus contains the determinants of hereditary qualities. In reproduction, the male sex-cell, which is scarcely more than a minute mass of chromatin provided with a thin coat of protoplasm and a motile organ, fuses with the egg, and the nuclei of the two cells unite to form a double body, which contains equal contributions of chromatin from the two parental organisms. This gives the physical basis for paternal inheritance as well as for maternal inheritance, and it shows why they may be of the same or equivalent degree. When, now, the egg divides, at the first and later cleavages, the chromatin masses or chromosomes contained in the double nucleus are split lengthwise and the twin portions separate to go into the nuclei of the daughter-cells. As the same process seems to hold for all the later divisions of the cleavage-cells whose products are destined to be the various tissue elements of the adult body, it follows that all tissue-cells would contain chromatin determinants derived equally from the male and female parents. As of course only the germ-cells of an adult organism pass on to form later generations, and as their content of chromatin is derived not from the sister organs of the body, but from the original fertilized egg, there is a direct stream of the germ plasm which flows continuously from the germ-cell to germ-cell through succeeding generations. It would seem, therefore, that the various organic systems are, so to speak, sister products in embryonic origin. The reproductive organs are not produced by the other parts of the body, but their cells are the direct descendants of the common starting-point namely, the egg. As the cells of the reproductive organs are the only ones that pass over and into the next and later generations, it will be evident, in the first place, that the germ plasm of their nuclei is the only essential substance that connects parent and offspring. This stream of germ plasm passes on in direct continuity through successive generations—from egg to the complete adult, including its own germ-cells, through these to the next adult, with its germ-cells, and so on and on as long as the species exists. It does not flow circuitously from egg to adult and then to new germ-cells, but it is direct and continuous, and apparently it cannot pick up any of the body-changes of an acquired nature. Now we see why individual acquisitions are not transmitted. The hereditary stream of germ plasm is already constituted before an animal uses its parts in adult life; we cannot see how alterations in the structure of mature body parts through use and adjustment to the environment can be introduced into it to become new qualities of the species.

It must be clear, I am sure, that this theory supplements natural selection, for it describes the physical basis of inheritance, it demonstrates the efficiency of congenital or germ-plasmal factors of variation in contrast with the Lamarckian factors, and finally in the way that in the view of Weismann it accounts for the origin of variations as the result of the commingling of two differing parental streams of germ plasm.

At first, for many reasons, Weismann's theories did not meet with general acceptance, but during recent years there has been a marked return to many of his positions, mainly as the result of further cytological discoveries, and of the formulation of Mendel's Law and of De Vries's mutation theory. The first-named law was propounded by Gregor Mendel on the basis of extensive experiments upon plants conducted during many years, 1860 and later, in the obscurity of his monastery garden at Altbrünn, in Austria. It was rescued from oblivion by De Vries, who found it buried in a mass of literature and brought it to light when he published his renowned Mutation Theory in 1901. Mendelian phenomena of inheritance, confirmed and extended by numerous workers with plants and animals, prove that in many cases portions of the streams of germ plasm that combine to form the hereditary content of organisms may retain their individuality during embryonic and later development, and that they may emerge in their original purity when the germ-cells destined to form a later generation undergo the preparatory processes of maturation. They demonstrate also the apparent chance nature of the phenomena of inheritance. To my mind the most striking and significant result in this field is the demonstration that a particular chromosome or chromatin mass determines a particular character of an adult organism, which is quite a different matter from the reference of all the hereditary characters to the chromatin as a whole. Wilson and others have brought forward convincing proof that the complex character of sex in insects actually resides in or is determined by particular and definite masses of this wonderful physical basis of inheritance.

Mendel's principles also account in the most remarkable way for many previously obscure phenomena, like reversion, or a case where a child resembles its grandparent more than it does either of its parents; such phenomena are due, so to speak, to the rise to the surface of a hidden stream of germ plasm that had flowed for one or many generations beneath its accompanying currents. I believe that the law is replacing more and more the laws of Galton and Pearson, formulated as statistical summaries of certain phenomena of human inheritance taken en masse. According to Galton's celebrated law of ancestral inheritance, the qualities of any organism are determined to the extent of a certain fraction by its two parents taken together as a "mid-parent," that a smaller definite fraction is contributed by the grandparents taken together as a mid-grandparent, and so on to earlier generations. But Mendel's Law has far greater definiteness, it explains more accurately the cases of alternative inheritance, and it may be shown to hold for blended and mosaic inheritance as well.

De Vries's new "mutation theory" is clearly not an alternative but a complementary theory to natural selection, the Weismannian and Mendelian theories. Like these last, it emphasizes the importance of the congenital hereditary qualities contained in the germ plasm, though unlike the Darwinian doctrine it shows that sometimes new forms may arise by sudden leaps and not necessarily by the slow and gradual accumulation of slight modifications or fluctuations. The mutants like any other variants must present themselves before the jury of environmental circumstances, which passes judgment upon their condition of adaptation, and they, too, must abide by the verdict that means life or death.

From what has been said of these post-Darwinian discoveries, the Lamarckian doctrine, which teaches that acquired non-congenital characters are transmitted, seems to be ruled out. I would not lead you to believe that the matter is settled. I would say only that the non-transmission of racial mutilations, negative breeding experiments upon mutilated rats and mice, the results of further study of supposedly transmitted immunity to poisons—that all these have led zoölogists to render the verdict of "not proved." The future may bring to light positive evidence, and cases like Brown-Séquard's guinea-pigs, and results like those of MacDougal with plants, and of Tower with beetles, may lead us to alter the opinion stated. But as it stands now most investigators hold that there are strong general grounds for disbelief in the principle, and also that it lacks experimental proof.

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The explanation of natural evolution given by Darwinism and the principles of Weismann, Mendel, and De Vries, still fails to solve the mystery completely, and appeal has been made to other agencies, even to teleology and to "unknown" and "unknowable" causes as well as to circumstantial factors. A combination of Lamarckian and Darwinian factors has been proposed by Osborn, Baldwin, and Lloyd Morgan, in the theory of organic selection. The theory of orthogenesis propounded by Naegeli and Eimer, now gaining much ground, holds that evolution takes place in direct lines of progressive modification, and is not the result of apparent chance. Of these and similar theories, all we can say is that if they are true, they are not so well substantiated as the ones we have reviewed at greater length.

The task of experimental zoölogy is to work more extensively and deeply upon inheritance and variation, combining the methods and results of cellular biology, biometrics, and experimental breeding. We may safely predict that great advances will be made during the next few years in analyzing the method of evolution; and that a few decades hence men will look back to the present time as a period of transition like the era of reawakened interest and renewed investigation that followed the appearance of the "Origin of Species." For the present, we can justly say "that evolution, so far as it is understood, is a real and natural process."