The Principles of Biology Vol. I/Chapter II.7

§ 75. Having, in the last chapter but one, concluded what constitutes an individual, and having, in the last chapter, contemplated the histological process which initiates a new individual, we are in a position to deal with the multiplication of individuals. For this, the title Genesis is here chosen as being the most comprehensive title—the least specialized in its meaning. By some biologists Generation has been used to signify one method of multiplication, and Reproduction to signify another method; and each of these words has been thus rendered in some degree unfit to signify multiplication in general.

Here the reader is indirectly introduced to the fact that the production of new organisms is carried on in fundamentally unlike ways. Up to quite recent times it was believed, even by naturalists, that all the various processes of multiplication observable in different kinds of organisms, have one essential character in common: it was supposed that in every species the successive generations are alike. It has now been proved, however, that in many plants and in numerous animals, the successive generations are not alike; that from one generation there proceeds another whose members differ more or less in structure from their parents; that these produce others like themselves, or like their parents, or like neither; but that eventually, the original form re-appears. Instead of there being, as in the cases most familiar to us, a constant recurrence of the same form, there is a cyclical recurrence of the same form. These two distinct processes of multiplication, may be aptly termed homogenesis and heterogenesis. Under these heads let us consider them.

There are two kinds of homogenesis, the simplest of them, probably once universal but now exceptional, being that in which there is no other form of multiplication than one resulting from perpetual spontaneous fission. The rise of distinct sexes was doubtless a step in evolution, and before it took place the formation of new individuals could have arisen only by division of the old, either into two or into many. At present this process survives, so far as appears, among Bacteria, certain Algæ, and sundry Protozoa; though it is possible that a rarely-occurring conjugation has in these cases not yet been observed. It is a probable conclusion, however, that in the Bacteria at any rate, the once universal mode of multiplication still survives as an exceptional mode. But now passing over these cases, we have to note that the kind of genesis (once supposed to be the sole kind), in which the successive generations are alike, is sexual genesis, or, as it has been otherwise called—gamogenesis. In every species which multiplies by this kind of homogenesis, each generation consists of males and females; and from the fertilized germs they produce the next generation of similar males and females arises: the only needful qualification of this statement being that in many Protophyta and Protozoa the conjugating cells or protoplasts are not distinguishable in character. This mode of propagation has the further trait, that each fertilized germ usually gives rise to but one individual—the product of development is organized round one axis and not round several axes, Homogenesis in contrast with heterogenesis as exhibited in species which display distinct sexuality, has also the characteristic that each new individual begins as an egg detached from the maternal tissues, instead of being a portion of protoplasm continuous with them, and that its development proceeds independently. This development may be carried on either internally or externally; whence results the division into the oviparous and the viviparous. The oviparous kind is that in which the fertilized germ is extruded from the parent before it has undergone any considerable development. The viviparous kind is that in which development is considerably advanced, or almost completed, before extrusion takes place. This distinction is, however, not a sharply-defined one: there are transitions between the oviparous and the viviparous processes. In ovo-viviparous genesis there is an internal incubation; and though the young are in this case finally extruded from the parent in the shape of eggs, they do not leave the parent's body until after they have assumed something like the parental form. Looking around, we find that homogenesis is universal among the Vertebrata. Every vertebrate animal arises from a fertilized germ, and unites into its single individuality the whole product of this fertilized germ. In the mammals or highest Vertebrata, this homogenesis is in every case viviparous; in birds it is uniformly oviparous; and in reptiles and fishes it is always essentially oviparous, though there are cases of the kind above referred to, in which viviparity is simulated. Passing to the Invertebrata, we find oviparous homogenesis universal among the Arachnida (except the Scorpions, which are ovo-viviparous); universal among the higher Crustacea, but not among the lower; extremely general, though not universal, among Insects; and universal among the higher Mollusca though not among the lower. Along with extreme inferiority among animals, we find homogenesis to be the exception rather than the rule; and in the vegetal kingdom there appear to be no cases, except among the Algæ and a few aberrant parasites like the Rafflesiaceæ, in which the centre or axis which arises from a fertilized germ becomes the immediate producer of fertilized germs.

In propagation characterized by unlikeness of the successive generations, there is asexual genesis with occasionally-recurring sexual genesis; in other words—agamogenesis interrupted more or less frequently by gamogenesis. If we set out with a generation of perfect males and females, then, from their ova arise individuals which are neither males nor females, but which produce the next generation from buds. By this method of multiplication many individuals originate from a single fertilized germ. The product of development is organized round more than one centre or axis. The simplest form of heterogenesis is that seen in most uniaxial plants. If, as we find ourselves obliged to do, we regard each separate shoot or axis of growth as a distinct individual, homogenesis is seen in those which have absolutely terminal flowers; but in all other uniaxial plants, the successive individuals are not represented by the series A, A, A, A, &c., but they are represented by the series A, B, A, B, A, B, &c. For in the majority of plants which were classed as uniaxial ( § 50), and which may be conveniently so distinguished from other plants, the axis which shoots up from the seed, and substantially constitutes the plant, does not itself flower but gives lateral origin to flowering axes. Though in ordinary uniaxial plants the fructifying apparatus appears to be at the end of the primary, vertical axis; yet dissection shows that, morphologically considered, each fructifying axis is an offspring from the primary axis. There arises from the seed a sexless individual, from which spring by gemmation individuals having reproductive organs; and from these there result fertilized germs or seeds that give rise to sexless individuals. That is to say, gamogenesis and agamogenesis alternate: the peculiarity being that the sexual individuals arise from the sexless ones by continuous development. The Salpæ show us an allied form of heterogenesis in the animal kingdom. Individuals developed from fertilized ova, instead of themselves producing fertilized ova, produce, by gemmation, strings of individuals from which fertilized ova again originate. In multiaxial plants, we have a succession of generations represented by the series A, B, B, B, &c., A, B, B, B, &c. Supposing A to be a flowering axis or sexual individual, then, from any fertilized germ it casts off, there grows up a sexless individual, B; from this there bud-out other sexless individuals, B, and so on for generations more or less numerous, until at length, from some of these sexless individuals, there bud-out seed-bearing individuals of the original form A. Branched herbs, shrubs, and trees, exhibit this form of heterogenesis: the successive generations of sexless individuals thus produced being, in most cases, continuously developed, or aggregated into a compound individual, but being in some cases discontinuously developed. Among animals a kind of heterogenesis represented by the same succession of letters, occurs in such compound polypes as the Sertularia, and in those of the Hydrozoa which assume alternately the polypoid form and the form of the Medusa. The chief differences presented by these groups arise from the fact that the successive generations of sexless individuals produced by budding, are in some cases continuously developed, and in others discontinuously developed; and from the fact that, in some cases, the sexual individuals give off their fertilized germs while still growing on the parent-polypedom, but in other cases not until after leaving the parent-polypedom and undergoing further development. Where, as in all the foregoing kinds of agamogenesis, the new individuals bud out, not from any specialized reproductive organs but from unspecialized parts of the parent, the process has been named, by Prof. Owen, metagenesis. In most instances the individuals thus produced grow from the outsides of the parents—the metagenesis is external. But there is also a kind of metagenesis which we may distinguish as internal. Certain entozoa of the genus Distoma exhibit it. From the egg of a Distoma there results a rudely-formed creature known as a sporocyst and from this a redia. Gradually, as this divides and buds, the greater part of the inner substance is transformed into young animals called Cercariæ (which are the larvæ of Distomata); until at length it becomes little more than a living sac full of living offspring. In the Distoma pacifica, the brood of young animals thus arising by internal gemmation are not Cercariæ, but are like their parent: themselves becoming the producers of Cercariæ, after the same manner, at a subsequent period. So that now the succession of forms is represented by the series A, B, A, B, &c., now by the series A, B, B, A, B, B, &c., and now by A, B, B, C, A. Both cases, however, exemplify internal metagenesis in contrast with the several kinds of external metagenesis described above. That agamogenesis which is carried on in a reproductive organ—either an ovarium or the homologue of one—has been called, by Prof. Owen, parthenogenesis. It is the process familiarly exemplified in the Aphides. Here, from the fertilized eggs laid by perfect females there grow up imperfect females, in the ovaria of which are developed ova that though unfertilized, rapidly assume the organization of other imperfect females, and are born viviparously. From this second generation of imperfect females, there by-and-by arises, in the same manner, a third generation of the same kind; and so on for many generations: the series being thus symbolized by the letters A, B, B, B, B, B, &c., A. Respecting this kind of heterogenesis it should be added that, in animals as in plants, the number of generations of sexless individuals produced before the re-appearance of sexual ones, is indefinite; both in the sense that in the same species it may go on to a greater or less extent according to circumstances, and in the sense that among the generations of individuals proceeding from the same fertilized germ, a recurrence of sexual individuals takes place earlier in some of the diverging lines of multiplication than in others. In trees we see that on some branches flower-bearing axes arise while other branches are still producing only leaf-bearing axes; and in the successive generations of Aphides a parallel fact has been observed. Lastly has to be set down that kind of heterogenesis in which, along with gamogenesis, there occurs a form of agamogenesis exactly like it, save in the absence of fecundation. This is called true parthenogenesis—reproduction carried on by virgin mothers which are in all respects like other mothers. Among silk-worm-moths this parthenogenesis is exceptional rather than ordinary. Usually the eggs of these insects are fertilized; but if they are not they are still laid, and some of them produce larvæ. In certain Lepidoptera, however, of the groups Psychidæ and Tineidæ, parthenogenesis appears to be a normal process—indeed, so far as is known, the only process; for of some species the males have never been found.

A general conception of the relations among the different modes of Genesis, thus briefly described, will be best given by the following tabular statement.

This, like all other classifications of such phenomena, presents anomalies. It may be justly objected that the processes here grouped under the head agamogenesis, are the same as those before grouped under the head of discontinuous development ( § 50): thus making development and genesis partially coincident. Doubtless it seems awkward that what are from one point of view considered as structural changes are from another point of view considered as modes of multiplication. There is, however, nothing for us but a choice of imperfections. We cannot by any logical dichotomies accurately express relations which, in Nature, graduate into one another insensibly. Neither the above, nor any other scheme, can do more than give an approximate idea of the truth.

§ 76. Genesis under every form is a process of negative or positive disintegration; and is thus essentially opposed to that process of integration which is the primary process in individual evolution. Negative disintegration occurs in those cases where, as among the compound Hydrozoa, there is a continuous development of new individuals by budding from the bodies of older individuals; and where the older individuals are thus prevented from growing to a greater size, or reaching a higher degree of integration. Positive disintegration occurs in those forms of agamogenesis where the production of new individuals is discontinuous, as well as in all cases of gamogenesis. The degrees of disintegration are various. At the one extreme the parent organism is completely broken up, or dissolved into new individuals; and at the other extreme each new individual forms but a small deduction from the parent organism. Protozoa and Protophyta show us that form of disintegration called spontaneous fission: two or more individuals being produced by the splitting-up of the original one. The Volvox and the Hydrodictyon are plants which, having developed broods within themselves, give them exit by bursting; and among animals the one lately referred to which arises from the Distoma egg, entirely loses its individuality in the individualities of the numerous Distoma-larvæ with which it becomes filled. Speaking generally, the degree of disintegration becomes less marked as we approach the higher organic forms. Plants of superior types throw off from themselves, whether by gamogenesis or agamogenesis, parts that are relatively small; and among superior animals there is no case in which the parent individuality is habitually lost in the production of new individuals. To the last, however, there is of necessity a greater or less disintegration. The seeds and pollen-grains of a flowering plant are disintegrated portions of tissue; as are also the ova and spermatozoa of animals. And whether the fertilized germs carry away from their parents small or large quantities of nutriment, these quantities in all cases involve further negative or positive disintegrations of the parents.

Except in spore-producing plants, new individuals which result from agamogenesis usually do not separate from the parent-individuals until they have undergone considerable development, if not complete development. The agamogenetic offspring of those lowest organisms which develop centrally, do not, of course, pass beyond central structure; but the agamogenetic offspring of organisms which develop axially, commonly assume an axial structure before they become independent. The vegetal kingdom shows us this in the advanced organization of detached bulbils, and of buds that root themselves before separating. Of animals, the Hydrozoa, the Trematoda, and the Salpæ, present us with different kinds of agamogenesis, in all of which the new individuals are organized to a considerable extent before being cast off. This rule is not without exceptions, however. The statoblasts of the Plumatella (which play the part of winter eggs), developed in an unspecialized part of the body, furnish a case of metagenesis in which centres of development, instead of axes, are detached; and in the above-described parthenogenesis of moths and bees, such centres are detached from an ovarium.

When produced by gamogenesis, the new individuals become (in a morphological sense) independent of the parents while still in the shape of centres of development, rather than axes of development; and this even where the reverse is apparently the case. The fertilized germs of those inferior plants which are central, or multicentral, in their development, are of course thrown off as centres; and the same is usually the case even in those which are uniaxial or multiaxial. In the higher plants, of the two elements that go to the formation of the fertilized germ, the pollen-cell is absolutely separated from the parent-plant under the shape of a centre, and the egg-cell, though not absolutely separated from the parent, is still no longer subordinate to the organizing forces of the parent. So that when, after the egg-cell has been fertilized by matter from the pollen-tube, the development commences, it proceeds without parental control: the new individual, though remaining physically united with the old individual, becomes structurally and functionally separate: the old individual doing no more than supply materials. Throughout the animal kingdom, the new individuals produced by gamogenesis are obviously separated in the shape of centres of development wherever the reproduction is oviparous: the only conspicuous variation being in the quantity of nutritive matter bequeathed by the parent at the time of separation. And though, where the reproduction is viviparous, the process appears to be different, and in one sense is so, yet, intrinsically, it is the same. For in these cases the new individual really detaches itself from the parent while still only a centre of development; but instead of being finally cast off in this state it is re-attached, and supplied with nutriment until it assumes a more or less complete axial structure.

§ 77. As we have lately seen, the essential act in gamogenesis is the union of two cell-nuclei, produced in the great majority of cases by different parent organisms. Nearly always the containing cells, often called gametes, are unlike: the sperm-cell being the male product, and the germ-cell the female. But among some Protozoa and many of the lower Algæ and Fungi, the uniting cells show no differentiation. Sexuality is only nascent.

There are very many modes and modifications of modes in which these cells are produced; very many modes and modifications of modes by which they are brought into contact; and very many modes and modifications of modes by which the resulting fertilized germs have secured to them the fit conditions for their development. But passing over these divergent and re-divergent kinds of sexual multiplication, which it would take too much space here to specify, the one universal trait is this coalescence of a detached portion of one organism with a more or less detached portion of another.

Such simple Algæ as the Desmidieæ, which are sometimes called unicellular plants, show us a coalescence, not of detached portions of two organisms, but of two entire organisms: the entire contents of the individuals uniting to form the germ-mass. Where, as among the Confervoideæ, we have aggregated cells whose individualities are scarcely at all subordinate to that of the aggregate, the gamogenetic act is often effected by the union "of separate motile protoplasmic masses produced by the division of the contents of any cell of the aggregate. These free-swimming masses of protoplasm, which are quite similar to (but generally smaller than) the agamogenetic 'zoospores' of the same plants, and to the free-swimming individuals of many Protophyta, are apparently the primitive type of gametes (conjugating cells); but it is noteworthy that such a gamete nearly always unites with one derived from another cell or from another individual. The same fact holds with regard to the gametes of the Protophytes themselves, which are formed in the same way from the single cell of the mother individual. In the higher types of Confervoideæ, and in Vaucheria, we find these equivalent, free-swimming, gametes replaced by sexually differentiated sperm- and germ-cells, in some cases arising in different organs set apart for their production, and essentially representing those found in the higher plants. Transitional forms, intermediate between these and the cases where equivalent gametes are formed from any cell of the plant are also known."

Recent investigations concerning the conjugation of Protozoa have shown that there is not, as was at one time thought, a fusion of two individualities, but a fusion of parts of their nuclei. The macro-nucleus having disappeared, and the micro-nucleus having broken up into portions, each individual receives from the other one of these portions, which becomes fused with its own nuclear matter. So that even in these humble forms, where there is no differentiation of sexes, the union is not between elements that have arisen in the same individual but between those which have arisen in different individuals: the parts being in this case alike.

The marvellous phenomena initiated by the meeting of sperm-cell and germ-cell, or rather of their nuclei, naturally suggest the conception of some quite special and peculiar properties possessed by these cells. It seems obvious that this mysterious power which they display of originating a new and complex organism, distinguishes them in the broadest way from portions of organic substance in general. Nevertheless, the more we study the evidence the more are we led towards the conclusion that these cells are not fundamentally different from other cells. The first fact which points to this conclusion is the fact recently dwelt upon ( § 63), that in many plants and inferior animals, a small fragment of tissue which is but little differentiated, is capable of developing into an organism like that from which it was taken. This implies that the component units of tissues have inherent powers of arranging themselves into the forms of the organisms which originated them. And if in these component units, which we distinguished as physiological, such powers exist,—if, under fit conditions, and when not much specialized, they manifest such powers in a way as marked as that in which the contents of sperm-cells and germ-cells manifest them; then, it becomes clear that the properties of sperm-cells and germ-cells are not so peculiar as we are apt to assume. Again, the organs emitting sperm-cells and germ-cells have none of the specialities of structure which might be looked for, did sperm-cells and germ-cells need endowing with properties unlike those of all other organic agents. On the contrary, these reproductive centres proceed from tissues characterized by their low organization. In plants, for example, it is not appendages that have acquired considerable structure which produce the fructifying particles: these arise at the extremities of the axes where the degree of structure is the least. The cells out of which come the egg and the pollen-grains, are formed from undifferentiated tissue in the interior of the ovule and of the stamen. Among many inferior animals devoid of special reproductive organs, such as the Hydra, the ova and spermatozoa originate from the interstitial cells of the ectoderm, which lie among the bases of the functional cells—have not been differentiated for function; and in the Medusæ, according to Weismann, they arise in the homologous layer, save where the medusoid form remains attached, and then they arise in the endoderm and migrate to the ectoderm: lack of specialization being in all cases implied. Then in the higher animals these same generative agents appear to be merely modified epithelium-cells—cells not remarkable for their complexity of structure but rather for their simplicity. If, by way of demurrer to this view, it be asked why other epithelium-cells do not exhibit like properties; there are two replies. The first is that other epithelium-cells are usually so far changed to fit them to their special functions that they are unfitted for assuming the reproductive function. The second is that in some cases, where they are but little specialized, they do exhibit the like properties: not, indeed, by uniting with other cells to produce new germs but by producing new germs without such union. I learn from Dr. Hooker that the Begonia phyllomaniaca habitually develops young plants from the scales of its stem and leaves—nay, that many young plants are developed by a single scale. The epidermal cells composing one of these scales swell, here and there, into large globular cells; form chlorophyll in their interiors; shoot out rudimentary axes; and then, by spontaneous constrictions, cut themselves off; drop to the ground; and grow into Begonias. Moreover, in a succulent English plant, the Malaxis paludosa, a like process occurs: the self-detached cells being, in this case, produced by the surfaces of the leaves. Thus, there is no warrant for the assumption that sperm-cells and germ-cells possess powers fundamentally unlike those of other cells. The inference to which the facts point, is, that they differ from the rest mainly in not having undergone functional adaptations. They are cells which have departed but little from the original and most general type: such specializations as some of them exhibit in the shape of locomotive appliances, being interpretable as extrinsic modifications which have reference to nothing beyond certain mechanical requirements. Sundry facts tend likewise to show that there does not exist the profound distinction we are apt to assume between the male and female reproductive elements. In the common polype sperm-cells and germ-cells are developed in the same layer of indifferent tissue; and in Tethya, one of the sponges, Prof. Huxley has observed that they occur mingled together in the general parenchyma. The pollen-grains and embryo-cells of plants arise in adjacent parts of the meristematic tissue of the flower-bud; and from the description of a monstrosity in the Passion-flower, recently given by Mr. Salter to the Linnæan Society, it appears both that ovules may, in their general structure, graduate into anthers, and that they may produce pollen in their interiors. Moreover, among the lower Algæ, which show the beginning of sexual differentiation, the smaller gametes, which we must regard as incipient sperm-cells, are sometimes able to fuse inter se, and give rise to a zygote which will produce a new plant. All which evidence is in perfect harmony with the foregoing conclusion; since, if sperm-cells and germ-cells have natures not essentially unlike those of unspecialized cells in general, their natures cannot be essentially unlike each other.

The next general fact to be noted is that these cells whose union constitutes the essential act of gamogenesis, are cells in which the developmental changes have come to a close—cells which are incapable of further evolution. Though they are not, as many cells are, unfitted for growth and metamorphosis by being highly specialized, yet they have lost the power of growth and metamorphosis. They have severally reached a state of equilibrium. And while the internal balance of forces prevents a continuance of constructive changes, it is readily overthrown by external destructive forces. For it almost uniformly happens that sperm-cells and germ-cells which are not brought in contact disappear. In a plant, the egg-cell, if not fertilized, is absorbed or dissipated, while the ovule aborts; and the unimpregnated ovum eventually decomposes: save, indeed, in those types in which parthenogenesis is a part of the normal cycle.

Such being the characters of these cells, and such being their fates if kept apart, we have now to observe what happens when they are united. In plants the extremity of the elongated pollen-cell applies itself to the surface of the embryo-sac, and one of its nuclei having, with some protoplasm, passed into the egg-cell, there becomes fused with the nucleus of the egg-cell. Similarly in animals, the spermatozoon passes through the limiting membrane of the ovum, and a mixture takes place between the substance of its nucleus and the substance of the nucleus of the ovum. But the important fact which it chiefly concerns us to notice, is that on the union of these reproductive elements there begins, either at once or on the return of favourable conditions, a new series of developmental changes. The state of equilibrium at which each had arrived is destroyed by their mutual influence, and the constructive changes, which had come to a close, recommence. A process of cell-multiplication is set up; and the resulting cells presently begin to aggregate into the rudiment of a new organism.

Thus, passing over the variable concomitants of gamogenesis, and confining our attention to what is constant in it, we see:—that there is habitually, if not universally, a fusion of two portions of organic substance which are either themselves distinct individuals, or are thrown off by distinct individuals; that these portions of organic substance, which are severally distinguished by their low degree of specialization, have arrived at states of structural quiescence or equilibrium; that if they are not united this equilibrium ends in dissolution; but that by the mixture of them this equilibrium is destroyed and a new evolution initiated.

§ 78. What are the conditions under which Genesis takes place? How does it happen that some organisms multiply by homogenesis and others by heterogenesis? Why is it that where agamogenesis prevails it is usually from time to time interrupted by gamogenesis? A survey of the facts discloses certain correlations which, if not universal, are too general to be without significance.

Where multiplication is carried on by heterogenesis we find, in numerous cases, that agamogenesis continues as long as the forces which result in growth are greatly in excess of the antagonist forces. Conversely, we find that the recurrence of gamogenesis takes place when the conditions are no longer so favourable to growth. In like manner where there is homogenetic multiplication, new individuals are usually not formed while the preceding individuals are still rapidly growing—that is, while the forces producing growth exceed the opposing forces to a great extent; but the formation of new individuals begins when nutrition is nearly equalled by expenditure. A few out of the many facts which seem to warrant these inductions must suffice.

The relation in plants between fructification and innutrition (or rather, between fructification and such diminished nutrition as makes growth relatively slow) was long ago asserted by a German biologist—Wolff, I am told. Since meeting with this assertion I have examined into the facts for myself. The result has been a conviction, strengthened by every inquiry, that some such relation exists. Uniaxial plants begin to produce their lateral, flowering axes, only after the main axis has developed the great mass of its leaves, and is showing its diminished nutrition by smaller leaves, or shorter internodes, or both. In multiaxial plants two, three, or more generations of leaf-bearing axes, or sexless individuals, are produced before any seed-bearing individuals show themselves. When, after this first stage of rapid growth and agamogenetic multiplication, some gamogenetic individuals arise, they do so where the nutrition is least;—not on the main axis, or on secondary axes, or even on tertiary axes, but on axes that are the most removed from the channels which supply nutriment. Again, a flowering axis is commonly less bulky than the others: either much shorter or, if long, much thinner. And further, it is an axis of which the terminal internodes are undeveloped: the foliar organs, which instead of becoming leaves become sepals, and petals, and stamens, follow each other in close succession, instead of being separated by portions of the still-growing axis. Another group of evidences meets us when we observe the variations of fruit-bearing which accompany variations of nutrition in the plant regarded as a whole. Besides finding, as above, that gamogenesis commences only when growth has been checked by extension of the remoter parts to some distance from the roots, we find that gamogenesis is induced at an earlier stage than usual by checking the nutrition. Trees are made to fruit while still quite small by cutting their roots or putting them into pots; and luxuriant branches which have had the flow of sap into them diminished, by what gardeners call "ringing," begin to produce flower-shoots instead of leaf-shoots. Moreover, it is to be remarked that trees which, by flowering early in the year, seem to show a direct relation between gamogenesis and increasing nutrition, really do the reverse; for in such trees the flower-buds are formed in the autumn. That structure which determines these buds into sexual individuals is given when the nutrition is declining. Conversely, very high nutrition in plants prevents, or arrests, gamogenesis. It is notorious that unusual richness of soil, or too large a quantity of manure, results in a continuous production of leaf-bearing or sexless shoots; and a like result happens when the cutting down of a tree, or of a large part of it, is followed by the sending out of new shoots: these, supplied with excess of sap, are luxuriant and sexless. Besides being prevented from producing sexual individuals by excessive nutrition, plants are, by excessive nutrition, made to change the sexual individuals they were about to produce, into sexless ones. This arrest of gamogenesis may be seen in various stages. The familiar instance of flowers made barren by the transformation of their stamens into petals, shows us the lowest degree of this reversed metamorphosis. Where the petals and stamens are partially changed into green leaves, the return towards the agamogenetic structure is more marked; and it is still more marked when, as occasionally happens in luxuriantly-growing plants, new flowering axes, and even leaf-bearing axes, grow out of the centres of flowers.Among various examples I have observed, the most remarkable were among Foxgloves, growing in great numbers and of large size, in a wood between Whatstandwell Bridge and Crich, in Derbyshire. In one case the lowest flower on the stem contained, in place of a pistil, a shoot or spike of flower-buds, similar in structure to the embryo-buds of the main spike. I counted seventeen buds on it; of which the first had three stamens, but was otherwise normal; the second had three; the third, four; the fourth, four; &c. Another plant, having more varied monstrosities, evinced excess of nutrition with equal clearness. The following are the notes I took of its structure:—1st, or lowest flower on the stem, very large; calyx containing eight divisions, one partly transformed into a corolla, and another transformed into a small bud with bract (this bud consisted of a five-cleft calyx, four sessile anthers, a pistil, and a rudimentary corolla); the corolla of the main flower, which was complete, contained six stamens, three of them bearing anthers, two others being flattened and coloured, and one rudimentary; there was no pistil but, in place of it, a large bud, consisting of a three-cleft calyx of which two divisions were tinted at the ends, an imperfect corolla marked internally with the usual purple spots and hairs, three anthers sessile on this mal-formed corolla, a pistil, a seed vessel with ovules, and, growing to it, another bud of which the structure was indistinct. 2nd flower, large; calyx of seven divisions, one being transformed into a bud with bract, but much smaller than the other; corolla large but cleft along the top; six stamens with anthers, pistil, and seed-vessel. 3rd flower, large; six-cleft calyx, cleft corolla, with six stamens, pistil, and seed-vessel, with a second pistil half unfolded at its apex. 4th flower, large; divided along the top, six stamens. 5th flower, large; corolla divided into three parts, six stamens. 6th flower, large; corolla cleft, calyx six cleft, the rest of the flower normal. 7th, and all succeeding flowers, normal.

While this chapter is under revision, another noteworthy illustration has been furnished to me by a wall-trained pear tree which was covered in the spring by luxuriant "foreright" shoots. As I learned from the gardener, it was pruned just as the fruit was setting. A large excess of sap was thus thrown into other branches, with the result that in a number of them the young pears were made monstrous by reversion. In some cases, instead of the dried up sepals at the top of the pear, there were produced good sized leaves; and in other cases the seed-bearing core of the pear was transformed into a growth which protruded through the top of the pear in the shape of a new shoot. The anatomical structure of the sexual axis affords corroborative evidence: giving the impression, as it does, of an aborted sexless axis. Besides lacking those internodes which the leaf-bearing axis commonly possesses, the flowering axis differs by the absence of rudimentary lateral axes. In a leaf-bearing shoot the axil of every leaf usually contains a small bud, which may or may not develop into a lateral shoot; but though the petals of a flower are homologous with leaves, they do not bear homologous buds at their bases. Ordinarily, too, the foliar appendages of sexual axes are much smaller than those of sexless ones—the stamens and pistils especially, which are the last formed, being extremely dwarfed; and it may be that the absence of chlorophyll from the parts of fructification is a fact of like meaning. Moreover, the formation of the seed-vessel appears to be a direct consequence of arrested nutrition. If a gloved-finger be taken to represent a growing shoot, (the finger standing for the pith of the shoot and the glove for the peripheral layers of meristem and young tissue, in which the process of growth takes place); and if it be supposed that there is a diminished supply of material for growth; then, it seems a fair inference that growth will first cease at the apex of the axis, represented by the end of the glove-finger; and supposing growth to continue in those parts of the peripheral layers of young tissue that are nearer to the supply of nutriment, their further longitudinal extension will lead to the formation of a cavity at the extremity of the shoot, like that which results in a glove-finger when the finger is partially withdrawn and the glove sticks to its end. Whence it seems, both that this introversion of the apical meristem may be considered as due to failing nutrition, and that the ovules growing from its introverted surface (which would have been its outer surface but for the defective nutrition) are extremely aborted homologues of external appendages: both they and the pollen-grains being either morphologically or literally quite terminal, and the last showing by their dehiscence the exhaustion of the organizing power.

Those kinds of animals which multiply by heterogenesis, present us with a parallel relation between the recurrence of gamogenesis and the recurrence of conditions checking rapid growth: at least, this is shown where experiments have thrown light on the connexion of cause and effect; namely, among the Aphides. These creatures, hatched from eggs in the spring, multiply by agamogenesis, which in this case is parthenogenesis, throughout the summer. When the weather becomes cold and plants no longer afford abundant sap, perfect males and females are produced; and from gamogenesis result fertilized ova. But beyond this evidence we have much more conclusive evidence. For it has been shown, both that the rapidity of the agamogenesis is proportionate to the warmth and nutrition, and that if the temperature and supply of food be artificially maintained, the agamogenesis continues through the winter. Nay more—it not only, under these conditions, continues through one winter, but it has been known to continue for four successive years: some forty or fifty sexless generations being thus produced. And those who have investigated the matter see no reason to doubt the indefinite continuance of this agamogenetic multiplication, so long as the external requirements are duly met. Evidence of another kind, complicated by special influences, is furnished by the heterogenesis of the Daphnia—a small crustacean commonly known as the Water-flea, which inhabits ponds and ditches. From the nature of its habitat this little creature is exposed to very variable conditions. Besides being frozen in winter, the small bodies of water in which it lives are often unduly heated by the summer Sun, or dried up by continued drought. The circumstances favourable to the Daphnia's life and growth, being thus liable to interruptions which, in our climate, have a regular irregularity of recurrence; we may, in conformity with the hypothesis, expect to find both that the gamogenesis recurs along with declining physical prosperity and that its recurrence is very variable. I use the expression "declining physical prosperity" advisedly; since "declining nutrition," as measured by supply of food, does not cover all the conditions. This is shown by the experiments of Weismann (abstracted for me by Mr. Cunningham) who found that in various Daphnideæ which bring forth resting eggs, sexual and asexual reproduction go on simultaneously, as well as separately, in the spring and summer: these variable results being adapted to variable conditions. For not only are these creatures liable to die from lack of food, from the winter's cold, and from the drying up of their ditches, &c., as well as from the over-heating of them, but during this period of over-heating they are liable to die from that deoxygenation of the water which heat causes. Manifestly the favourable and unfavourable conditions recurring in combinations that are rarely twice alike, cannot be met by any regularly recurring form of heterogenesis; and it is interesting to see how survival of the fittest has established a mixed form. In the spring, as well as in the autumn, there is in some cases a formation of resting or winter eggs; and evidently these provide against the killing off of the whole population by summer drought. Meanwhile, by ordinary males and females there is a production of summer eggs adapted to meet the incident of drying up by drought and subsequent re-supply of water. And all along successive generations of parthenogenetic females effect a rapid multiplication as long as conditions permit. Since life and growth are impeded or arrested not by lack of food only, but by other unfavourable conditions, we may understand how change in one or more of these may set up one or other form of genesis, and how the mixture of them may cause a mixed mode of multiplication which, originally initiated by external causes, becomes by inheritance and selection a trait of the species. And then in proof that external causes initiate these peculiarities, we have the fact that in certain Daphnideæ "which live in places where existence and parthenogenesis are possible throughout the year, the sexual period has disappeared:" there are no males.

Passing now to animals which multiply by homogenesis—animals in which the whole product of a fertilized germ aggregates round a single centre or axis instead of round many centres or axes—we see, as before, that so long as the conditions allow rapid increase in the mass of this germ-product, the formation of new individuals by gamogenesis does not take place. Only when growth is declining in relative rate, do perfect sperm-cells and germ-cells begin to appear; and the fullest activity of the reproductive function arises as growth ceases: speaking generally, at least; for though this relation is tolerably definite in the highest orders of animals which multiply by gamogenesis, it is less definite in the lower orders. This admission does not militate against the hypothesis, as it seems to do; for the indefiniteness of the relation occurs where the limit of growth is comparatively indefinite. We saw ( § 46) that among active, hot-blooded creatures, such as mammals and birds, the inevitable balancing of assimilation by expenditure establishes, for each species, an almost uniform adult size; and among creatures of these kinds (birds especially, in which this restrictive effect of expenditure is most conspicuous), the connexion between cessation of growth and commencement of reproduction is distinct. But we also saw ( § 46) that where, as in the Crocodile and the Pike, the conditions and habits of life are such that expenditure does not overtake assimilation as size increases, there is no precise limit of growth; and in creatures thus circumstanced we may naturally look for a comparatively indeterminate relation between declining growth and commencing reproduction. There is, indeed, among fishes, at least one case which appears very anomalous. The male parr, or young of the male salmon, a fish of four or five inches in length, is said to produce milt. Having, at this early stage of its growth, not one-hundredth of the weight of a full-grown salmon, how does its production of milt consist with the alleged general law? The answer must be in great measure hypothetical. If the salmon is (as it appears to be in its young state) a species of fresh-water trout that has contracted the habit of annually migrating to the sea, where it finds a food on which it thrives—if the original size of this species was not much greater than that of the parr (which is nearly as large as some varieties of trout)—and if the limit of growth in the trout tribe is very indefinite, as we know it to be; then we may reasonably infer that the parr has nearly the adult form and size which this species of trout had before it acquired its migratory habit; and that this production of milt is, in such case, a concomitant of the incipient decline of growth naturally arising in the species when living under the conditions of the ancestral species. Should this be so, the immense subsequent growth of the parr into the salmon, consequent on a suddenly-increased facility in obtaining food, removes to a great distance the limit at which assimilation is balanced by expenditure; and has the effect, analogous to that produced in plants, of arresting the incipient reproductive process. A confirmation of this view may be drawn from the fact that when the parr, after its first migration to the sea, returns to fresh water, having increased in a few months from a couple of ounces to five or six pounds, it no longer shows any fitness for propagation: the grilse, or immature salmon, does not produce milt or spawn.

We conclude, then, that the products of a fertilized germ go on accumulating by simple growth, so long as the forces whence growth results are greatly in excess of the antagonist forces; but that when diminution of the one set of forces or increase of the other, causes a considerable decline in this excess and an approach towards equilibrium, fertilized germs are again produced. Whether the germ-product be organized round one axis or round the many axes that arise by agamogenesis, matters not. Whether, as in the higher animals, this approach to equilibrium results from that disproportionate increase of expenditure entailed by increase of size; or whether, as in most plants and many inferior animals, it results from absolute or relative decline of nutrition; matters not. In any case the recurrence of gamogenesis is associated with a decrease in the excess of tissue-producing power. We cannot say, indeed, that this decrease always results in gamogenesis: some organisms multiply for an indefinite period by agamogenesis only. The Weeping Willow, which has been propagated throughout Europe, does not seed in Europe; and yet, as the Weeping Willow, by its large size and the multiplication of generation upon generation of lateral axes, presents the same causes of local innutrition as other trees, we cannot ascribe the absence of sexual axes to the continued predominance of nutrition. Among animals, too, the anomalous case of the Tineidæ, a group of moths in which parthenogenetic multiplication goes on for generation after generation, seems to imply that gamogenesis does not necessarily result from an approximate balance of assimilation by expenditure. What we must say is that an approach towards equilibrium between the forces which cause growth and the forces which oppose growth, is the chief condition to the recurrence of gamogenesis; but that there appear to be other conditions, in the absence of which approach to equilibrium is not followed by gamogenesis.

§ 79. The above induction is an approximate answer to the question—When does gamogenesis recur? but not to the question which was propounded—Why does gamogenesis recur?—Why cannot multiplication be carried on in all cases, as it is in many cases, by agamogenesis? As already said, biologic science is not yet advanced enough to reply. Meanwhile, the evidence above brought together suggests a certain hypothetical answer.

Seeing, on the one hand, that gamogenesis recurs only in individuals which are approaching a state of organic equilibrium; and seeing, on the other hand, that the sperm-cells and germ-cells thrown off by such individuals are cells in which developmental changes have ended in quiescence, but in which, after their union, there arises a process of active cell-formation; we may suspect that the approach towards a state of general equilibrium in such gamogenetic individuals, is accompanied by an approach towards molecular equilibrium in them; and that the need for this union of sperm-cell and germ-cell is the need for overthrowing this equilibrium, and re-establishing active molecular change in the detached germ—a result probably effected by mixing the slightly different physiological units of slightly different individuals. The several arguments which support this view, cannot be satisfactorily set forth until after the topics of Heredity and Variation have been dealt with. Leaving it for the present, I propose hereafter to re-consider it in connexion with sundry others raised by the phenomena of Genesis.

But before ending the chapter, it may be well to note the relations between these different modes of multiplication, and the conditions of existence under which they are respectively habitual. While the explanation of the teleologist is untrue, it is often an obverse to the truth; for though, on the hypothesis of Evolution, it is clear that things are not arranged thus or thus for the securing of special ends, it is also clear that arrangements which do secure these special ends tend to establish themselves—are established by their fulfilment of these ends. Besides insuring a structural fitness between each kind of organism and its circumstances, the working of "natural selection" also insures a fitness between the mode and rate of multiplication of each kind of organism and its circumstances. We may, therefore, without any teleological implication, consider the fitness of homogenesis and heterogenesis to the needs of the different classes of organisms which exhibit them.

Heterogenesis prevails among organisms of which the food, though abundant compared with their expenditure, is dispersed in such a way that it cannot be appropriated in a wholesale manner. Protophyta, subsisting on diffused gases and decaying organic matter in a state of minute subdivision, and Protozoa, to which food comes in the shape of extremely small floating particles, are enabled, by their rapid agamogenetic multiplication, to obtain materials for growth better than they would do did they not thus continually divide and disperse in pursuit of it. The higher plants, having for nutriment the carbonic acid of the air and certain mineral components of the soil, show us modes of multiplication adapted to the fullest utilization of these substances. A herb with but little power of forming the woody fibre requisite to make a stem that can support wide-spreading branches, after producing a few sexless axes produces sexual ones; and maintains its race better, by the consequent early dispersion of seeds, than by a further production of sexless axes. But a tree, able to lift its successive generations of sexless axes high into the air, where each gets carbonic acid and light almost as freely as if it grew by itself, may with advantage go on budding-out sexless axes year after year; since it thereby increases its subsequent power of budding-out sexual axes. Meanwhile it may advantageously transform into seed-bearers those axes which, in consequence of their less direct access to materials absorbed by the roots, are failing in their nutrition; for it thus throws off from a point at which sustenance is deficient, a migrating group of germs that may find sustenance elsewhere. The heterogenesis displayed by animals of the Cœlenterate type has evidently a like utility. A polype, feeding on minute annelids and crustaceans which, flitting through the water, come in contact with its tentacles, and limited to that quantity of prey which chance brings within its grasp, buds out young polypes which, either as a colony or as dispersed individuals, spread their tentacles through a larger space of water than the parent alone can; and by producing them, the parent better insures the continuance of its species than it would do if it went on slowly growing until its nutrition was nearly balanced by its waste, and then multiplied by gamogenesis. Similarly with the Aphis. Living on sap sucked from tender shoots and leaves, and able thus to take in but a very small quantity in a given time, this creature's race is more likely to be preserved by a rapid asexual propagation of small individuals, which disperse themselves over a wide area of nutrition, than it would be did the individual growth continue so as to produce large individuals multiplying sexually. And then when autumnal cold and diminishing supply of sap put a check to growth, the recurrence of gamogenesis, or production of fertilized ova which remain dormant through the winter, is more favourable to the preservation of the race than would be a further continuance of agamogenesis. On the other hand, among the higher animals living on food which, though dispersed, is more or less aggregated into large masses, this alternation of gamic and agamic reproduction ceases to be useful. The development of the germ-product into a single organism of considerable bulk, is in many cases a condition without which these large masses of nutriment could not be appropriated; and here the formation of many individuals instead of one would be fatal. But we still see the beneficial results of the general law—the postponement of gamogenesis until the rate of growth begins to decline. For so long as the rate of growth continues rapid, there is proof that the organism gets food with facility—that expenditure does not seriously check accumulation; and that the size reached is as yet not disadvantageous: or rather, indeed, that it is advantageous. But when the rate of growth is much decreased by the increase of expenditure—when the excess of assimilative power is diminishing so fast as to indicate its approaching disappearance—it becomes needful, for the maintenance of the species, that this excess shall be turned to the production of new individuals; since, did growth continue until there was a complete balancing of assimilation and expenditure, the production of new individuals would be either impossible or fatal to the parent. And it is clear that "natural selection" will continually tend to determine the period at which gamogenesis commences, in such a way as most favours the maintenance of the race.

Here, too, may fitly be pointed out the fact that, by "natural selection," there will in every case be produced the most advantageous proportion of males and females. If the conditions of life render numerical inequality of the sexes beneficial to the species, in respect either of the number of the offspring or the character of the offspring; then, those varieties of the species which approach more than other varieties towards this beneficial degree of inequality, will be apt to supplant other varieties. And conversely, where equality in the number of males and females is beneficial, the equilibrium will be maintained by the dying out of such varieties as produce offspring among which the sexes are not balanced.

—Such alterations of statement in this chapter as have been made necessary by the advance of biological knowledge since 1864 have not, I think, tended to invalidate its main theses, but have tended to verify them. Some explanations to be here added may remove remaining difficulties.

Certain types, which are transitional between Protozoa and Metazoa, exhibit under its simplest form the relation between self-maintenance and race-maintenance—the integration primarily effecting the one and the disintegration primarily effecting the other. Among the Mycetozoa a number of amœba-like individuals aggregate into what is called a plasmodium; and while, in some orders, they become fused into a mass of protoplasm through which their nuclei are dispersed, in other orders (Sorophora) they retain their individualities and simply form a coherent aggregate. These last, presumably the earliest in order of evolution, remain united so long as the plasmodium, having a small power of locomotion, furthers the general nutrition; but when this is impeded by drought or cold, there arise spores. Each spore contains an amœboid individual; and this, escaping when favourable conditions return, establishes by fission and by union with others like itself a new colony or plasmodium. Reduced to its lowest terms, we here see the antagonism between that growth of the coherent mass of units which accompanies its physical prosperity, and that incoherence and dispersion of the units which follows unfavourable conditions and arrest of growth, and which presently initiates new plasmodia.

This antagonism, seen in these incipient Metazoa which show us none of that organization characterizing the Metazoa in general, is everywhere in more or less disguised forms exhibited by them—must necessarily be so if growth of the individual is a process of integration while formation of new individuals is a process of disintegration. And, primarily, it is an implication that whatever furthers the one impedes the other.

But now while recognizing the truth that nutrition and innutrition (using these words to cover not supply of nutriment only but the presence of other influences favourable or unfavourable to the vital processes) primarily determine the alternations of these; we have also to recognize the truth that from the beginning survival of the fittest has been shaping the forms and effects of their antagonism. By inheritance a physiological habit which modifies the form of the antagonism in a way favourable to the species, will become established. Especially will this be the case where the lives of the individuals have become relatively definite and where special organs have been evolved for casting off reproductive centres. The resulting physiological rhythm may in such cases become so pronounced as greatly to obscure the primitive relation. Among plants we see this in the fact that those which have been transferred from one habitat to another having widely different seasons, long continue their original time of flowering, though it is inappropriate to the new circumstances—the reproductive periodicity has become organic. Similarly in each species of higher animal, development of the reproductive organs and maturation of reproductive cells take place at a settled age, whether the conditions have been favourable or unfavourable to physical prosperity. The established constitutional tendency, adapted to the needs of the species, over-rides the constitutional needs of the individual.

Even here, however, the primitive antagonism, though greatly obscured, occasionally shows itself. Instance the fact that in plants where gamogenesis is commencing a sudden access of nutrition will cause resumption of agamogenesis; and I suspect that an illustration may be found among human beings in the earlier establishment of the reproductive function among the ill-fed poor than among the well-fed rich.

One other qualification has to be added. In plants and animals which have become so definitely constituted that at an approximately fixed stage, the proclivity towards the production of new individuals becomes pronounced, it naturally happens that good nutrition aids it. Surplus nutriment being turned into the reproductive channel, the reproduction is efficient in proportion as the surplus is great. Hence the fact that in fruit trees which have reached the flowering stage, manuring has the effect that though it does not increase the quantity of blossoms it increases the quantity of fruit; and hence the fact that well-fed and easy-living races of men are prolific.