LAWS OF TRANSMISSION BY INHERITANCE. ADAPTATION AND NUTRITION.
Distinction between Conservative and Progressive Transmission by Inheritance.—Laws of Conservative Transmission: Transmission of Inherited Characters.—Uninterrupted or Continuous Transmission.—Interrupted or Latent Transmission.—Alternation of Generations.—Relapse.—Degeneracy.—Sexual Transmission.—Secondary Sexual Characters.—Mixed or Amphigonous Transmission.—Hybrids.—Abridged or Simplified Transmission.—Laws of Progressive Inheritance: Transmission of Acquired Characters.—Adapted or Acquired Transmission.—Fixed or Established Transmission.—Homochronous Transmission (Identity in Epoch).—Homotopic Transmission (Identity in Part).—Adaptation and Mutability.—Connection between Adaptation and Nutrition.—Distinction between Indirect and Direct Adaptation.
In the last chapter we considered Transmission by Inheritance, one of the two universal vital activities of organisms, Adaptation and Inheritance, which by their interaction produce the different species of organisms, and we have endeavoured to trace this very mysterious vital activity to a more general physiological function of organisms, namely, to Propagation. This latter in its turn, like other vital phenomena of animals and plants, depends on physical and chemical relations. It is true they appear at times exceedingly complicated, but can nevertheless in reality be traced to simple mechanical causes—that is, to the relations of attraction and repulsion in the particles or molecules—in fact, to the motional phenomena of matter.
Now, before we turn our attention to the second function, the phenomenon of Adaptation or Mutability, which counteracts the Transmission by Inheritance, it seems appropriate first to cast one more glance at the various manifestations of Heredity, which we may perhaps even now denominate the “laws of transmission by inheritance.” Unfortunately, up to the present time very little has been done for this most important subject, either in zoology or in botany, and almost all we know of the different laws of inheritance is confined to the experiences of gardeners and farmers. It is not therefore to be wondered at, that on the whole these exceedingly interesting and important phenomena have not been investigated with desirable scientific accuracy, or reduced to the form of scientific laws. Accordingly, what I shall relate of the different laws of transmission are only some preliminary fragments taken out of the infinitely rich store which lies open to our inquiry.
We may first divide all the different phenomena of inheritance into two groups, which we may distinguish as the transmission of inherited characters, and the transmission of acquired characters; and we may call the former the conservative transmission, and the latter the progressive transmission by inheritance. This distinction depends upon the exceedingly important fact that the individuals of every species of animals and plants can transmit to their descendants, not only those qualities which they themselves have inherited from their ancestors, but also the peculiar, individual qualities which they have acquired during their own life. The latter are transmitted by progressive, the former by conservative inheritance. We have now first to examine the phenomena of conservative inheritance, that is, the transmission of such qualities as the organism has already received from its parents or ancestors. (Gen. Morph. ii. 180.)
Among the phenomena of conservative inheritance we are first struck by that which is its most general law, and which we may term the law of uninterrupted or continuous transmission. It is so universal among the higher animals and plants, that the uninitiated might overestimate its action and consider it as the only normal law of transmission by inheritance. This law simply consists in the fact that among most species of animals and plants, every generation is, on the whole, like the preceding—that the parents are as like the grandparents as they are like the children. “Like produces like,” as is commonly said, but more accurately “similar things produce similar things.” For, in reality, the descendants of every organism are never absolutely equal in all points, but only similar in a greater or less degree. This law is so generally known, that I need not give any examples of it.
The law of interrupted or latent transmission by inheritance, which might also be termed alternating transmission, is in a measure opposed to the preceding law. This important law appears principally active among many lower animals and plants, and manifests itself in contrast to the former in the fact that the offspring are not like their parents, but very dissimilar, and that only the third or a later generation becomes similar to the first. The grandchildren are like the grandparents, but quite unlike the parents. This is a remarkable phenomenon, and, as is well known, occurs also very frequently, though in a less degree, in human families. Every one of my readers doubtless knows some members of a family who, in this or that peculiarity, much more resemble the grandfather or grandmother than the father or mother. Sometimes it lies in bodily peculiarities, for example, features of face, colour of hair, size of body—sometimes in mental qualities, for example, temperament, energy, understanding—which are transmitted in this manner. This fact may be observed in domestic animals as well as in the case of man. Among the domestic animals most liable to vary—as the dog, horse, and ox—breeders very frequently find that the product by breeding resembles the grandparents far more than it does its own parental organism. If we express this general law and the succession of generations by the letters of the alphabet, then A = C = E, whilst B = D = F, and so on.
This very remarkable fact appears in a more striking way in the lower animals and plants than in the higher, and especially in the well-known phenomenon of alternation of generations (metagenesis). Here we very frequently find—for example, among the Planarian worms, sea-squirts or Tunicates, Zoophytes, and also among ferns and mosses—that the organic individual in the first place produces, by propagation, a form completely different from the parental form, and that only the descendants of this generation, again, become like the first. This regular change of generation was discovered by the poet Chamisso, on his voyage round the world in 1819, among the Salpæ, cylindrical tunicates, transparent like glass, which float on the surface of the sea. Here the larger generation, the individuals of which live isolated and possess an eye of the form of a horse-shoe, produce in a non-sexual manner (by the formation of buds) a completely different and smaller generation. The individuals of this second smaller generation live united in chains and possess a cone-shaped eye. Every individual of such a chain produces, in a sexual manner (hermaphrodite) again, a non-sexual solitary form of the first and larger generation. Among the Salpæ, therefore, it is always the first, third, and fifth generation, and in like manner the second, fourth, and sixth generations, that are entirely like one another. However, it is not always only one, but in other cases a number of generations, which are thus leapt over; so that the first generation resembles the fourth and seventh, the second resembles the fifth and eighth, the third resembles the sixth and ninth, and so on. Three different generations alternate with one another; for example, among the neat little sea-buoys (Doliolum), small tunicates closely related to the Salpæ. In this case it is A = D = G, further, B = E = H, and C = F = I. Among the plant-lice (Aphides), each sexual generation is followed by a succession of from eight to ten or twelve non-sexual generations, which are like one another, but differ from the sexual generations. Then, again, a sexual generation reappears like the one long before vanished.
If we further follow this remarkable law of latent or interrupted inheritance, and take into consideration all the phenomena appertaining to it, we may comprise under it also the well-known phenomena of reversion. By the term “reversion” or “atavism” we understand the remarkable fact known to all breeders of animals, that occasionally single and individual animals assume a form which has not existed for many generations, but belongs to a generation which has long since disappeared. One of the most remarkable instances of this kind is the fact that in some horses there sometimes appear singular dark stripes, similar to those of the zebra, quagga, and other wild species of African horses. Domestic horses of the most different races and of all colours sometimes show such dark stripes; for example, a stripe along the back, a stripe across the shoulders, and the like. The sudden appearance of these stripes can only be explained by the supposition that it is the effect of a latent transmission, a relapse into the ancient original form, which has long since vanished, and was once common to all species of horses; the original form, undoubtedly, was originally striped like the zebras, quaggas, etc. In like manner, certain qualities in other domestic animals sometimes appear quite suddenly, which once marked their wild ancestors, now long since extinct. In plants, also, such a relapse can be observed very frequently. All my readers probably know the wild yellow toad-flax (Linaria vulgaris), a plant very common in our fields and hedges. Its dragon-mouthed yellow flower contains two long and two short stamens. But sometimes there appears a single blossom (Peloria) which is funnel-shaped, and quite regularly composed of five individual and equal sections, with five corresponding stamens. This Peloria can only be explained as a relapse into the long since extinct and very ancient common form of all those plants which, like the toad-flax, possess dragon-mouthed, two-lipped flowers, with two long and two short stamens. The original form, like the Peloria, possessed a regular five-spurred blossom, with five equal stamens, which only later and by degrees have become unequal (compare p. [17]). All such relapses are to be brought under the law of interrupted or latent transmission, although the number of intervening generations may be enormous.
When cultivated plants or domestic animals become wild, when they are withdrawn from the conditions of cultivated life, they experience changes which appear not only as adaptations to their new mode of life, but partially also as relapses into the ancient original form out of which the cultivated forms have been developed. Thus the different kinds of cabbage, which are exceedingly different in form, may be led back to the original form, by allowing them to grow wild. In like manner, dogs, horses, heifers, etc., when growing wild, often revert more or less to a long extinct generation. An immensely long succession of generations may pass away before this power of latent transmission becomes extinguished.
A third law of conservative transmission may be called the law of sexual transmission, according to which each sex transmits to the descendants of the same sex peculiarities which are not inherited by the descendants of the other sex. The so-called secondary sexual characters, which in many respects are of extraordinary interest, everywhere furnish numerous examples of this law. Subordinate or secondary sexual characters are those peculiarities of one of the two sexes which are not directly connected with the sexual organs themselves; such characters, which exclusively belong to the male sex, are, for example, the antlers of the stag, the mane of the lion, and the spur of the cock. The human beard, an ornament commonly denied to the female sex, belongs to the same class. Similar characteristics by which the female sex is alone distinguished are, for example, the developed breasts, with the lactatory glands of female mammals and the pouch of the female opossum. The bodily size, also, and complexion, differs in female animals of many species from that of the male. All these secondary sexual qualities, like the sexual organs themselves, are transmitted by the male organism only to the male, not to the female, and vice versâ. Contrary facts are rare exceptions to the rule.
A fourth law of transmission, which has here to be mentioned, in a certain sense contradicts the last, and limits it, viz., the law of mixed or mutual (amphigonous) transmission. This law tells us that every organic individual produced in a sexual way receives qualities from both parents, from the father as well as from the mother. This fact, that personal qualities of each of the two sexes are transmitted to both male and female descendants, is very important, Goethe mentions it of himself, in the beautiful lines—
“Von Vater hab ich die Statur, des Lebens ernstes Führen Von Mütterchen die Frohnatur und Lust zu fabuliren.” “From my father I have my stature and the serious tenour of my life, From my mother a joyous nature and a turn for poetizing.”
This phenomenon, I suppose, is so well-known to all, that I need not here enter upon it. It is according to the different portions of their character which father and mother transmit to their children, that the individual differences among brothers and sisters are chiefly determined.
The very important and interesting phenomenon of hybridism also belongs to this law of mixed or amphigonous transmission. It alone, when rightly estimated, is quite sufficient to refute the prevailing dogma of the constancy of species. Plants, as well as animals, belonging to quite different species, may sexually mingle with one another and produce descendants which in many cases can again propagate themselves, and that indeed either (more frequently) by mingling with one of the two parental species, or (more rarely) by pure in-breeding, hybrid mixing with hybrid. The latter is well established, for example, in the hybrids of hares and rabbits (Lepus Darwinii, p. 147). The hybrids of a horse and a donkey, two different species of the same genus (Equus), are well known. These hybrids differ according as the father or the mother belongs to the one or the other species—the horse or the donkey. The mule produced by a mare and a he-donkey has qualities quite different from those of the jinny (Hinnus), the hybrid of a horse and she-donkey. In both cases the hybrid produced by the crossing of two different species is a mixed form, which receives qualities from both parents; but the qualities of the hybrid are different, according to the form of the crossing. In like manner, mulattoes produced by a European and a negress show a different mixture of characters from the hybrids produced by a negro with a European female. In these phenomena of hybrid-breeding, as well as in the other laws of transmission previously mentioned, we are as yet unable to show the acting causes in detail; but no naturalist doubts the fact that the causes are in all cases purely mechanical and dependent upon the nature of organic matter itself. If we possessed more delicate means of investigation than our rude organs of sense and auxilliary instruments, we should be able to discover those causes, and to trace them to the chemical and physical properties of matter.
Among the phenomena of conservative transmission, we must now mention, as the fifth law, the law of abridged or simplified transmission. This law is very important in regard to embryology or ontogeny, that is in regard to the history of the development of organic individuals. Ontogeny, or the history of the development of individuals, as I have already mentioned in the first chapter (p. 10), and as I subsequently shall explain more minutely, is nothing but a short and quick repetition of Phylogeny dependent on the laws of transmission and adaptation—that is, a repetition of the palæontological history of development of the whole organic tribe, or phylum, to which the organism belongs. If, for example, we follow the individual development of a man, an ape, or any other higher mammal within the maternal body from the egg, we find that the fœtus or embryo arising out of the egg passes through a series of very different forms, which on the whole agrees with, or at least runs parallel to, a series of forms which is presented to us by the historical chain of ancestors of the higher mammals. Among these ancestors we may mention certain fishes, amphibians, marsupials, etc. But the parallelism or agreement of these two series of development is never quite complete; on the contrary, in ontogeny there are always gaps and leaps which indicate the omission of certain stages belonging to the phylogeny. Fritz Müller, in his excellent work, “Für Darwin,”[(16)] has clearly shown in the case of the Crustacea, or crabs, that “the historical record preserved in the individual history of development is gradually obscured, in proportion as development takes a more and more direct route from the egg to the complete animal.” This process of obscuring and shortening is determined by the law of abridged transmission, and I mention it here specially because it is of great importance for the understanding of embryology, and because it explains the fact, at first so strange, that the whole series of forms which our ancestors have passed through in their gradual development are no longer visible in the series of forms of our own individual development from the egg.
Opposed to the laws of the conservative transmission, hitherto discussed, are the phenomena of the transmission of the second series, that is, the laws of progressive transmission by inheritance. As already mentioned, they depend upon the fact that the organism transmits to its descendants not only those qualities which it has inherited from its own ancestors, but also a number of those individual qualities which it has acquired during its own lifetime. Adaptation is here seen to be connected with transmission by inheritance (Gen. Morph. ii. 186).
At the head of these important phenomena of progressive transmission, we may mention the law of adapted or acquired transmission. In reality it asserts nothing more than what I have said above, that in certain circumstances the organism is capable of transmitting to its descendants all the qualities which it has acquired during its own life by adaptation. This phenomenon, of course, shows itself most distinctly when the newly acquired peculiarity produces any considerable change in the inherited form. This is the case in the examples I mentioned in the preceding chapter as to transmission in general, in the case of the men with six fingers and toes, the porcupine men, copper beeches, weeping willows, etc. The transmission of acquired diseases, such as consumption, madness, and albinism, likewise form very striking examples. Albinoes are those individuals who are distinguished by the absence of colouring matter, or pigments, in the skin. They are of frequent occurrence among men, animals, and plants. In the case of animals of a definite dark colour, individuals are not unfrequently born which are entirely without colour, and in animals possessing eyes, this absence of pigment extends even to the eyes, so that the iris of the eye, which is commonly of a bright or intense colour, is colourless, but appears red, on account of the blood-vessels being seen through it. Among many animals, such as rabbits and mice, albinoes with white fur and red eyes are so much liked that they are propagated in great numbers as a special race. This would be impossible were it not for the law of the transmission of adaptations.
Which of the changes acquired by an organism are transmitted to its descendants, and which are not, cannot be determined à priori, and we are unfortunately not acquainted with the definite conditions under which the transmission takes place. We only know in a general way that certain acquired qualities are much more easily transmitted than others, for example, more easily than the mutilations caused by accidents. These latter are generally not transmitted by inheritance, otherwise the descendants of men who have lost their arms or legs would be born without the corresponding arm or leg; but here, also, exceptions occur, and a race of dogs without tails has been produced by consistently cutting off the tails of both sexes of the dog during several generations. A few years ago a case occurred on an estate near Jena, in which by a careless slamming of a stable door the tail of a bull was wrenched off, and the calves begotten by this bull were all born without a tail. This is certainly an exception; but it is very important to note the fact, that under certain unknown conditions such violent changes are transmitted in the same manner as many diseases.
In very many cases the change which is transmitted and preserved by adapted transmission is constitutional or inborn, as in the case of albinism mentioned before. The change then depends upon that form of adaptation which we call the indirect or potential. A very striking instance is furnished by the hornless cattle of Paraguay, in South America. A special race of oxen is there bred which is entirely without horns. It is descended from a single bull, which was born in 1770 of an ordinary pair of parents, and the absence of horns was the result of some unknown cause. All the descendants of this bull produced with a horned cow were entirely without horns. This quality was found advantageous, and by propagating the hornless cattle among one another, a hornless race was obtained, which at present has almost entirely supplanted the horned cattle in Paraguay. The case of the otter-sheep of North America forms a similar example. In the year 1791 a farmer, by name Seth Wright, lived in Massachusetts, in North America; in his normally formed flock of sheep a lamb was suddenly born with a surprisingly long body and very short and crooked legs. It was therefore unable to take any great leaps, and especially unable to leap across a hedge into a neighbour’s garden—a quality which seemed advantageous to the owner, as the territories were divided by hedges. It therefore occurred to him to transmit this quality to other sheep, and by crossing this ram with normally shaped ewes, he produced a whole race of sheep, all of which had the qualities of the father, short and crooked legs and a long body. None of them could leap across the hedges, and they therefore were much liked and propagated in Massachusetts.
A second law, which likewise belongs to the series of progressive transmissions, may be called the law of established or habitual transmission. It manifests itself in this, that qualities acquired by an organism during its individual life are the more certainly transmitted to its descendants the longer the causes of that change have been in action, and that this change becomes the more certainly the property of all subsequent generations the longer the cause of change acts upon these latter also. The quality newly acquired by adaptation or mutation must be established or constituted to a certain degree before we can calculate with any probability that it will be transmitted at all to the descendants. In this respect transmission resembles adaptation. The longer a newly acquired quality has been transmitted by inheritance, the more certainly will it be preserved in future generations. If, therefore, for example, a gardener by methodical treatment has produced a new kind of apple, he may calculate with the greater certainty upon preserving the desired peculiarity of this sort the longer he has transmitted the same by inheritance. The same is clearly shown in the transmission of diseases. The longer consumption or madness has been hereditary in a family the deeper is the root of the evil, and the more probable it is that all succeeding generations will suffer from it.
We may conclude the consideration of the phenomena of inheritance with the two very important laws of homotopic and contemporaneous transmission by inheritance. We understand by them the fact that changes acquired by an organism during its life, and transmitted to its descendants, appear in the same part of the body in which the parental organism was first affected by them, and that they also appear in the offspring at the same age as that at which they did so in the parent.
The law of contemporaneous or homochronous transmission, which Darwin calls the law of of “transmission in corresponding periods of life,” can be shown very clearly in the transmission of diseases, especially of such as are recognized as very destructive, on account of their hereditary character. They generally appear in the organism of the child at the time corresponding with that in which the parental organism contracted the disease. Hereditary diseases of the lungs, liver, teeth, brain, skin, etc., usually appear in the descendants at the same period, or a little earlier than they showed themselves in the parental organism, or were contracted by it. The calf gets its horns at the same period of life as its parents did. In like manner the young stag receives its antlers at the same period of life in which they appeared in its father or grandfather. In every one of the different sorts of vine the grapes ripen at the same time as they did in the case of their progenitors. It is well known that the time of ripening varies greatly in the different sorts; but as all are descended from a single species, this variation has been acquired by the progenitors of the several sorts, and has then been transmitted by inheritance.
The law of homotopic transmission, which is most closely connected with the last mentioned law, and which might be called the law of transmission in corresponding parts of the body, may also be very distinctly recognized in pathological cases of inheritance. Large moles, for example, or accumulations of pigment in several parts of the skin, tumours also, often appear during many generations, not only at the same period of life, but also in the same part of the skin. Excessive development of fat in certain parts of the body is likewise transmitted by inheritance. Above all, it is to be noted that numerous examples of this, as well as of the preceding law, may be found everywhere in the study of embryology. Both the law of homochronous and homotopic transmission are fundamental laws of embryology, or ontogeny. For these laws explain the remarkable fact that the different successive forms of individual development in all generations of one and the same species always appear in the same order of succession, and that the variations of the body always take place in the same parts. This apparently simple and self-evident phenomenon is nevertheless exceedingly wonderful and curious; we cannot explain its real causes, but may confidently assert that they are due to the direct transmission of the organic matter from the parental organism to that of the offspring, as we have seen above in the case of the process of transmission in general, by a consideration of the details of the various modes of reproduction.
Having thus, then, considered the most important laws of Inheritance, we now turn to the second series of phenomena bearing on natural selection, viz., to those of Adaptation or Variation. These phenomena, taken as a whole, stand in a certain opposition to the phenomena of Inheritance, and the difficulty which arises in examining them consists mainly in the two sets of phenomena being so completely intercrossed and interwoven. We are but seldom able to say with certainty—of the variations of form which occur before our eyes—how much is owing to Inheritance, and how much to Adaptation. All characters of form, by which organisms are distinguished, are caused either by Inheritance or by Adaptation; but as both functions are continually interacting with each other, it is extremely difficult for the systematic inquirer to recognize the share belonging to each of the two functions in the special structure of individual forms. This is, at present, all the more difficult, because we are as yet scarcely aware of the immense importance of this fact, and because most naturalists have neglected the theory of Adaptation, as well as that of Inheritance. The laws of Inheritance, which we have just discussed, as well as the laws of Adaptation, which we shall consider directly, in reality form only a small portion of the phenomena existing in this domain, but which have not as yet been investigated; and since every one of these laws can interact with every other, it is clear that there is an infinite complication of physiological actions, which are at work in the construction of organisms.
But now, as to the phenomenon of variation or adaptation in general, we must, as in the case of inheritance, view it as a quite universal, physiological fundamental quality of all organisms, without exception—as a manifestation of life which cannot be separated from the idea of organism. Strictly speaking, we must here also, as in the case of inheritance, distinguish between Adaptation itself and Adaptability. By Adaptation (Adaptio), or Variation (Variatio), we understand the fact that the organism, in consequence of influences of the surrounding outer world, assumes certain new peculiarities in its vital activity, composition, and form which it has not inherited from its parents; these acquired individual qualities are opposed to those which have been inherited, or, in other words, those which have been transmitted to it from its parents or ancestors. On the other hand, we call Adaptability (Adaptabilitas), or Variability (Variabilitas), the capability inherent in all organisms to acquire such new qualities under the influence of the outer world. (Gen. Morph. ii. 191.)
The undeniable fact of organic adaptation or variation is universally known, and can be observed at every moment in thousands of phenomena surrounding us. But just because the phenomena of variation by external influences appear so self-evident, they have hitherto undergone scarcely any accurate scientific investigation. To them belong all the phenomena which we look upon as the results of contracting and giving up habits, of practice and giving up practices, or as the results of training, of education, of acclimatization, of gymnastics, etc. Many permanent variations brought about by causes producing disease, that is to say, many diseases, are nothing but dangerous adaptations of the organism to injurious conditions of life. In the case of cultivated plants and domestic animals, variation is so striking and powerful that the breeder of animals and the gardener found their whole mode of proceeding upon it, or rather upon the interaction between these phenomena and those of Inheritance. It is also well known to every one that animals and plants, in their wild state, are subject to variation. Every systematic treatise on a group of animals or plants, if it were to be quite complete and exhaustive, ought to mention in every individual species the number of variations which differ more or less from the prevailing or typical form of the species. Indeed, in every careful systematic special treatise one finds, in the case of most species, mention of a number of such variations, which are described sometimes as individual deviations, and sometimes as so-called races, varieties, degenerate species, or subordinate species, and which often differ exceedingly from the original species, solely in consequence of the adaptation of the organism to the external conditions of life.
If we now endeavour to fathom the general causes of these phenomena of Adaptation, we arrive at the conclusion that in reality they are as simple as the causes of the phenomena of Inheritance. We have shown that the nature of the process of propagation furnishes the real explanation of the facts of Transmission by Inheritance, that is, the transmission of parental matter to the body of the offspring; and in like manner we can show that the physiological function of nutrition, or change of substance, affords a general explanation of Adaptation or Variation. When I here point to “nutrition” as the fundamental cause of variation and adaptation, I take this word in its widest sense, and I understand by it the whole of the material changes which the organism undergoes in all its parts through the influences of the surrounding outer world. Nutrition thus comprises not only the reception of actual nutritive substances and the influence of different kinds of food, but also, for example, the action upon the organism of water and of the atmosphere, the influence of sunlight, of temperature, and of all those meteorological phenomena which are implied in the term “climate.” The indirect and direct influence of the nature of the soil and of the dwelling-place also belong to it; and further, the extremely important and varied influence which is exercised upon every animal and every plant by the surrounding organisms, friends and neighbours, enemies and robbers, parasites, etc. All these and many other very important influences, all of which more or less modify the organism in its material composition, must be taken into consideration in studying the change of substance which goes on in living things. Adaptation, accordingly, is the consequence of all those material variations which are produced in the change of substance of the organism by the external conditions of existence, or by the influences of the surrounding external world.
How very much every organism is dependent upon the whole of its external surroundings, and changed by their alteration, is, in a general way, well known to every one. Only think how much the human power of action is dependent upon the temperature of the air, or how much the disposition of our minds depends upon the colour of the sky. Accordingly as the sky is cloudless and sunny, or covered with large heavy clouds, our state of mind is cheerful or dull. How differently do we feel and think in a forest during a stormy winter night and during a bright summer day! All the different moods of our soul depend upon purely material changes of our brain, upon movements of molecular plasma, which are started through the medium of the senses by the different influences of light, warmth, moisture, etc. “We are a plaything to every pressure of the air.” No less important and deeply influential are the effects produced upon our mind and body by the different quality and quantity of food. Our mental activity, the activity of our understanding and of our imagination, is quite different accordingly as we have taken tea or coffee, wine or beer, before or during our work. Our moods, wishes, and feelings are quite different when we are hungry and when we are satisfied. The national character of Englishmen and Gauchos, in South America, who live principally on meat and food rich in nitrogen, is wholly different from that of the Irish, feeding on potatoes, and that of the Chinese, living on rice, both of whom take food deficient in nitrogen. The latter also form much more fat than the former. Here, as everywhere, the variations of the mind go hand in hand with the corresponding transformations of the body; both are produced by purely material causes. But all other organisms, in the same way as man, are varied and changed by the different influences of nutrition. It is well known that we can change in an arbitrary way the form, size, colour, etc., of our cultivated plants and domestic animals, by change of food; that, for example, we can take from or give to a plant definite qualities, accordingly as we expose it to a greater or less degree of sunlight and moisture. As these phenomena are generally widely known, and as we shall proceed presently to the consideration of the different laws of adaptation, we will not dwell here any longer on the general facts of variation.
As the different laws of transmission may be naturally divided into the two series of conservative and progressive transmission, so we may also distinguish between two series of the laws of adaptation, first, the series of laws of indirect, and secondly, the series of laws of direct adaptation. The latter may also be called the laws of actual, and the former the laws of potential, adaptation.
The first series, comprising the phenomena of indirect (potential) adaptation, has, on the whole, hitherto been little attended to, and Darwin has the merit of having directed special attention to this series of changes. It is somewhat difficult to place this subject clearly before the reader; I will endeavour to make it clear hereafter by examples. Speaking quite generally, indirect or potential adaptation consists in the fact that certain changes in the organism, effected by the influence of nutrition (in its widest sense) and of the external conditions of existence in general, show themselves not in the individual form of the respective organism, but in that of its descendants. Thus, especially in organisms propagating themselves in a sexual way, the reproductive system, or sexual apparatus, is often influenced by external causes (which little affect the rest of the organism), to such a degree that its descendants show a complete alteration of form. This can be seen very strikingly in artificially produced monstrosities. Monstrosities can be produced by subjecting the parental organism to certain extraordinary conditions of life, and, curiously enough, such an extraordinary condition of life does not produce a change of the organism itself, but a change in its descendants. This cannot be called transmission by inheritance, because it is not a quality existing in the parental organism that is transmitted by inheritance. It is, on the contrary, a change affecting the parental organism, but not perceptible in it, that appears in the peculiar formation of its descendants. It is only the impulse to this new formation which is transmitted in propagation through the egg of the mother or the sperm of the father. The new formation exists in the parental organism only as a possibility (potential); in the descendants it becomes a reality (actual).
As this very important and very general phenomenon had hitherto been entirely neglected, people were inclined to consider all the visible variations and transformations of organic forms as phenomena of adaptation of the second series, that is, as phenomena of direct or actual adaptation. The essence of this latter kind of adaptation consists in the fact that the change affecting the organism (through nutrition, etc.) shows itself immediately by some transformation, and does not only make itself apparent in the descendants. To this class belong all the well-known phenomena in which we can directly trace the transforming influence of climate, food, education, training, etc., in their effects upon the individual itself.
We have seen how the two series of phenomena of progressive and conservative transmission, in spite of their difference in principle, in many ways interfere with and modify each other, and in many ways co-operate with and cross each other. The same is the case, in a still higher degree, in the two series of phenomena of indirect and direct adaptation, which are opposed to each other and yet closely connected. Some naturalists, especially Darwin and Carl Vogt, ascribe to the indirect or potential adaptation by far the more important and almost exclusive influence. But the majority of naturalists have hitherto been inclined to take the opposite view, and to attribute the principal influence to direct or actual adaptation. I consider this controversy, in the mean while, as almost useless. It is but seldom that we are in a condition, in any individual case of variation, to judge how much of it belongs to direct and how much to indirect adaptation. We are, on the whole, still too little acquainted with these exceedingly important and intricate relations, and can only assert, in a general way, that the transformation of organic forms is to be ascribed either to direct adaptation alone, or to indirect adaptation alone, or lastly, to the co-operation of both direct and indirect adaptation.