In discussing Mendelian varieties we consider the manner in which two contrasting characters—one present in the male parent and one in the female—are transmitted to the offspring. The characters in question may be the tallness of the male parent and the contrasting shortness of the female; or the brown eyes of the male and the blue eyes of the female; or the brown skin of the female parent and the white skin of the male one. These characters may be inherited in two ways: either they may be blended or they may remain distinct in the offspring. The children of the brown mother and the white father are usually coloured in some tint intermediate between those of the parents. The mulatto hybrid is fertile with either of the parent races, and again the offspring may take a tint intermediate between those of the parents, and so on through a number of generations. But somewhere in this series the concealed or recessive brown colour may appear in all its completeness, showing that it has been present in the organisations of all the intervening generations. The progeny of a tall male parent and a short female parent are not, in general, intermediate in stature between the parents; some of them may be tall and others short. The children of a brown-eyed mother and a blue-eyed father do not usually have eyes in which the colours of the parental eyes are blended: they are blue-eyed or brown-eyed. The contrasting characters are spoken of as dominant and recessive: if tallness is transmitted to offspring, which may nevertheless produce dwarf offspring, the latter character is said to be recessive to tallness. The contrasting characters of the parents therefore remain distinct in the progeny, some of the latter exhibiting the one character and some the other; while it may happen that the one character or the other may be segregated, so that it only appears in, and is transmitted by, the offspring. There are numerical relationships between the numbers of the offspring in which the contrasting characters appear.

Obviously, tallness and dwarfness are not characters which differ in quality: they are different degrees of the same thing. Brown eyes and blue eyes are not necessarily different in quality, for we may think of the same kinds of pigment as being present in the iris but mixed in different proportions. But the terminology of this branch of biology appears to suggest that the contrasting characters are, each of them, something quite different from the other: there are “factors” for “tallness,” “dwarfness,” for blue eyes and brown eyes, and so on. These qualities are called “unit-characters,” and they are supposed to possess much the same individuality in the germ-plasm as the “radicles” of the chemist possess in a compound. Sodium chloride, for instance, is not a blend of sodium and chlorine: the two kinds of atoms do not fuse together but are held together merely. The analogy is, however, very imperfect, for in the chemical molecule the characters are not those of either of the constituents but something quite different, whereas in the Mendelian cross the characters remain distinct, but one of them is patent while the other is latent. In the molecule, however, the atoms are regarded by the chemist as lying beside each other in certain positions, and the Mendelian factors are also spoken of as if they lay side by side in the germ-plasm. This terminology is useful, perhaps necessary, in the work of investigation, but we must not forget that it symbolises, rather than describes, the results of experiment. If the factors are identified with certain morphological structures in the nuclei of the germ-cells, obviously all the objections that may be urged against the Weismannian hypothesis as an hypothesis of development apply also to the Mendelian hypotheses as descriptions of a physical process of the transmission of morphological characters.

It should clearly be understood what is implied in the construction of such a hypothesis. Certain processes are observed to take place when a somatic cell divides: these processes we have regarded as having for their object the exact division of all the parts of the cell into two halves. This process of somatic cell division is modified when a germ cell divides prior to maturation (the process fitting it to become fertilised). Then the cell nucleus divides into four daughter-nuclei. One of these remains in the cell substance which is to become the ovum, and the other three, each of them invested in a minimal quantity of cytoplasm, are eliminated as the “polar bodies.” Also the number of chromosomes in the mother-cell becomes halved, so that the mature ovum, or spermatozoon, possesses only one-half of the number of chromosomes which are present in the ordinary somatic cell. Now let the reader puzzle out for himself what may be meant by this behaviour of the germ cells, and he will certainly see that several interpretations are possible. But suppose that the chromatin consists of an incredibly large number of bodies differing in chemical structure from each other, and occupying definite positions with regard to each other; and suppose that there is a mechanism of unimaginable complexity in the cell capable of rejecting some of these chemically individualised parts, and of “assembling” or arranging the others in much the same way as an engineer assembles the parts of a dynamo when he completes the machine. Then we may regard the hypothetical discrete bodies which form the hypothetical nuclear architecture as the material carriers of Mendelian characters. It is strange that the correspondence of such a logically constructed mechanism with the effects which it would produce if it existed should be regarded as a proof that it does exist, yet biological speculation has actually made use of such an argument. “It seems exceedingly unlikely that a mechanism so exactly adapted to bring it” (the separation from each other of the Mendelian material “factors” of inheritance) “about should be found in every developing germ cell if it had no connection with the segregation of characters that is observed in experimental breeding.” Put quite plainly this argument is as follows: there is a certain segregation to be seen in experimental breeding, and certain processes may be observed to occur in the developing germ cell. Add to these processes many others logically conceivable, and add to the observed material structure of the cell another structure also logically conceivable. Then the assumed mechanism and structure is “exactly adapted” to produce the effects which are to be explained. Therefore the mechanism and structure do actually exist!

That which renders the son similar to the father—the specific organisation—is undoubtedly very stable, and it may persist in the face of a variable environment. But now and then the son differs from the father. The differences may be “accidental” and may not be transmitted further—then we have to deal with an unstable fluctuation; or the differences may be permanent—then we have to deal with a stable mutation. What “produces” a mutation? A change of the environment, it may be said: if so, the mutation is an active change or adaptation of the organism to a change in its surroundings, and this adaptation is a permanent one and is transmitted. Or the mutation may be a spontaneous change of functioning. If this disturbance of the stability of the organisation is general, if it affects all the characters of the organism, we have to deal with the establishment of a new elementary species. But if the disturbance affects only one, or a few characters, then we need not recognise that a new elementary species has come into existence. Men and women remain men and women (in their morphology), although some time or other among the brown eyes characteristic of a race blue eyes may have appeared. The result of the disturbance, in this case, has been to cause one, or a few, of the characters that fluctuate to surpass their limits of stability.

The idea of the elementary species is a clear and simple one. It is a group of organisms connected by ties of blood relationship: all have descended from one pair of ancestors. The individuals exhibit certain characters, all of which are variable. This variability is not cumulative; in generation after generation the individuals of the species display variations which fluctuate round the same mean values. Two or more elementary species may have had the same origin—a common ancestor or ancestors—but the organisms in one species exhibit characters which, although similar in nature to those of the other species, yet fluctuate about different mean values.

This is not the “species” of the systematic biologist. The Linnean or systematic species is a concept which is much more difficult to define: it is a concept indeed which has not any clear and definite meaning, in actual practice.

We often forget how very young the science of systematic biology is, and how intimately its progress has been dependent on that of human invention and industrial enterprise. Physics and mathematics might be studied in a monastic cell, but the study of systematic biology can only be carried on when we have ships and other means of travelling—the means, in short, of collecting the animals and plants inhabiting all the parts of the earth’s surface. Until a comparatively few years ago the fauna and flora of great tracts of land and sea were almost unknown: even now our knowledge of the life of many parts of the earth is scanty and inaccurate. Systematic biology has therefore had to collect and describe the organisms of the earth, and in so doing it has set up the Linnean species of plants and animals. These we may describe as, in the main, categories of morphological structures. The older and more familiar species are clearly defined in this respect: such are cats and dogs, rabbits, tigers, herrings, lobsters, oysters, and so on: the individuals in each of these categories are clearly marked out with respect to their morphology, and the limits of the categories are clearly defined. In all of them the specific organisation has attained a high degree of stability so that the individuals “breed true to type”; and it has also attained a high degree of specialisation, so that it does not fuse with other organisations.

Yet, in the majority of the systematic species of biology, this criterion of specific individuality—this recognition of the isolation of the species from other species—cannot be applied. Very many species have been described from a few specimens only, many from only one. How does a systematist recognise that an organism with which he is dealing has not already been classified? It differs from all other organisms most like it, that is, he cannot identify it with any known specific description. But the differences may be very small, and if he had a number of specimens of the species most nearly resembling it he might find that these differences were less than the limits of variation in this most closely allied species, and he would then relegate it to this category. But if he has to compare his specimen with the “type” one, that is, the only existing specimen on which the species of comparison was founded, the test would be unavailable. The question to be answered is this: are the difference or differences to be regarded as fluctuations, or are they of “specific rank”? Now certainly many systematists of great experience possess this power of judgment, though they might be embarrassed by having to state clearly what were the grounds on which their judgment was based. But on the other hand hosts of species have been made by workers who did not possess this quality of judgment; and even with the great systematists of biology confusion has originated. Slowly, very slowly, the organic world is becoming better known, and this confusion is disappearing.

The species, then, whether it is the systematic group of the biological systems, or the elementary species based on the study of variability and inheritance, is an intellectual construction: an artifice designed to facilitate our description of nature. This is clearly the case with the higher orders of groups in classifications: genera, families, orders, classes, and phyla express logical relationships, or describe in a hypothetical form our notions of an evolutionary process. But species, it may be said, have an actual reality: there are no genera in nature, only species. These categories of organisms really exist; they have individuality, a certain kind of organic unity, inasmuch as the individuals composing them have descended from a common ancestor. Yet just as much may be said of genera, families, and the other groupings. One species originates from another by a process of transmutation: a genus is a group of species which have all had a common origin; a family is a similarly related group of genera, and so on. The higher categories of biological science are intended to introduce order and simplification into the confusion and richness of nature as we observe it, but obviously the concept of the species has the same practical object. Must we then say that there are no species in nature, only individuals? If so, we are at once embarrassed by the difficulty of forming a clear notion of what is meant by organic individuality. Does it not indicate that life on the earth is really integral, and that our analysis of its forms—species, genera, families, and so on—are only convenient ways of dealing actively with all its richness?