A Mechanico-Physiological Theory of Organic Evolution Part 2

If two corresponding characters, one derived from the father, the other from the mother, come into conflict in s.e.xual reproduction, the one or the other, or even a third alternative characteristic, which heretofore was present as a latent determinant, may develop in the child. But also both parental characters may appear at once and in various combinations.

Whether the development follows in the one way or the other depends on the strength of the individual determinants, on the kind of their idioplasmic arrangement, and on their agreement with the nature of the newly formed idioplasm.

16. HEREDITY AND VARIATION.

If heredity and variation are defined according to the true nature of organisms, they are only apparent opposites. Since idioplasm alone is transmitted from one ontogeny to the next following, the phylogenetic development consists solely in the continual progress of the idioplasm and the whole genealogical tree from the primordial drop of plasma up to the organism of the present day (plant or animal) is, strictly speaking, nothing else than an individual consisting of idioplasm, which at each ontogeny forms a new individual body, corresponding to its advance.

In this idioplasmic individual the _automatic_ or _perfecting variation_ is always active, so that the idioplasm of a phylogenetic line always grows by propagation of the determinants contained within it, as a tree grows larger through its whole duration of life by branching. On the other hand the _adaptation variation_ caused by external stimuli is present only in those periods of the phylogenetic line in which the idioplasm, and together with this the individual, do not possess the obtainable maximum of adaptation to their environment for the time being. Both of these variations of the idioplasm take place so slowly that only after a long series of generations do the new determinants become capable of developing and revealing themselves in the trans.m.u.tation of visible characters.

Aside from the phylogenetic variations already named, which take place according to the measure of ontogenetic growth, the idioplasm undergoes, as a result of crossing, as well as in changes of the ontogeny, _gamogenic variations_ which may be designated as stationary, since in the mingling of s.e.xually different idioplasms there arise only new arrangements of determinants already present, but no new formation of determinants takes place. Hence in this way arise also new combinations of developmental characteristics.

As a result of external injurious influences, abnormal variations, or _pathological variations_, appear in the idioplasm. These consist of disturbances of equilibrium, which take place also without new formation of determinants. Thereby the determinants already present are caused to develop in abnormal relations, and mostly in reversions.

Apart from the inheritable variations of the idioplasm just enumerated, and the transformations of visible characters involved in it, the soma-plasm and the non-plasmic substances experience, by the influence of nutrition and climate, greater or less variations, which const.i.tute _nutrition varieties_, and since the idioplasm remains unaffected in general, last only so long as the causes which called them forth.[F]

[F] Nageli, like Weismann, arrives at the conclusion that acquired characters are not inherited. He was not content, however, to rest the generalization upon purely speculative grounds, but undertook the experimental demonstration. After seventeen years of work by himself and son, especially upon several species of Hieracium, he satisfied himself that his theory was true to the facts. We all know now how far he fell short of settling the question.--_Trans._

If we have in mind the inner nature of the organism, there is, properly speaking, no such specific phenomenon as heredity, since the phylogenetic line is a continuous idioplasmic individual. In this sense heredity is nothing more than the persistence of organized substance in a movement in which variations are automatically induced, or the necessary transition of one idioplasmic configuration into the next following. It is present, not only among plant and animal individuals which are ontogenetically separated, but also everywhere within these individuals, where individual parts (cells, organs) follow each other in time. Hereditary phenomena are those that necessarily pa.s.s over to following generations, and in general those that are located in the idioplasm, since non-idioplasmic substance can be hereditary only through a limited number of cell generations.

Variations and heredity are generally estimated, not according to the inner nature of the mature individuals, but according to their relation in successive generations, since heredity is a.s.sumed when the ontogenetic characters remain the same, and variation when previously latent characters become visible. But these phenomena belong to another department of science; they concern the possibility and reality of development of the idioplasmic determinants.

17. VARIETY, RACE, MODIFICATION.

From the multifarious variations of organisms proceed various categories of kins.h.i.+p. _Varieties_ arise by extremely slow changes in the idioplasm due to the perfecting process and adaptation. Since these are conditioned by the same causes, they follow in all individuals of the same variety in uniform manner. Varieties are uniform, entirely constant under the most various external conditions, in general cross only with difficulty with related varieties, are not varied by accidental crosses, and persist through geological periods. Varieties belong to feral nature rather than to culture; they can a.s.sume all possible modifications without injury to their specific characteristics, but can show no distinctions of races, for all beginnings of race formation are destroyed by free intercrossing. They differ from species only in that they are to be designated as more closely related species, or species as more remotely related varieties. Every other distinguis.h.i.+ng characteristic is wanting.

_Races_ arise from gamogenic or pathological variations of the idioplasm. In the former case they presuppose crossing between related varieties or species, in the latter case an increased sensibility and weakening of the idioplasm. Very often both causes co-operate, since crossing follows more easily when the idioplasm is weakened by hurtful influences and since the irritability and weakening of the idioplasm increases if crossing has preceded. Race formation begins in single individuals. Among several individuals it begins in various directions because the causes are different and hence may display a great multiformity. Races are distinguished by more or less abnormal characteristics; they arise quickly--often in a single generation--and present various degrees of stability. This stability is insured to some extent only by the strictest in-and-in breeding. All races disappear through crossing, likewise many races that have arisen from pathological variations disappear even in s.e.xual reproduction (in self-fecundation).

Races belong exclusively to cultivation, where they can develop and exist protected from free intercrossing.

While varieties and races arise by progressional or stationary variation of the idioplasm, _modifications_ are produced by such influences of nutrition and climate as act only on the soma-plasm and the non-plasmic substances, and hence do not give rise to inheritable characters in the organism. Modifications persist only so long as their causes, and under other environments immediately pa.s.s over into the modifications corresponding to them. The transition is completed in the lowest plants during a limited number of cell generations; in an individual of the higher plants on the same stem during the growth of a single year. Each variety and each race appears clothed in a definite modification, and can change it within a range peculiar to itself.[A]

[G] The distinctions which Nageli here erects are, of course, purely arbitrary, and his definitions are suitable for use only in his own thesis.--_Trans._

18. SOCIAL AND INDIVIDUAL ORIGIN OF SPECIES.

The species arises neither from the _nutrition variety_ nor from the _race_; it is always a more advanced variety, and hence species formation is identical with variety formation. Cause for variation and consequently for variety formation is always shown, either when, environment remaining the same, the automatic variation of the idioplasm has advanced so far that the ontogeny is raised to a higher grade of organization and division of labor, or when external stimuli act for a sufficiently long time in a manner not in harmony with the previous adaptation. Hence various varieties arise easily from a uniform kins.h.i.+p, when these are thrown among unlike external influences by local separation, because in the separated places on the one hand the automatic evolution proceeds with unequal rapidity, and on the other hand adaptation takes place unequally.

But in general different varieties arise socially from a uniform kins.h.i.+p. This is because the related individuals living together are unequally stimulated on account of the great inequality of external influences which may exist at the smallest distances; and also because with slight individual differences unlike reactions often follow upon the same external influences. If identically similar individuals are equally inclined to very different reactions toward the same stimulus, sometimes the direction of the first variation decides the character of the adaptation and therefore the nature of the variety, because the variation, when once begun, progresses unswervingly even under somewhat different circ.u.mstances.[H] Hence divergent variations are found growing together in all places, which variations have begun at different though neighboring points by transformation of the idioplasm and are soon intermingled on account of the easy dissemination of seed.

[H] It is interesting to compare this statement with Weismann's recent theory of Germinal Selection.--_Trans._

The social formation of varieties is not in general interrupted by crossing, a process which governs only the formation of races. It is confirmed according to experience by the universally recurring fact that several beginnings of the most closely related varieties appear together not only in the same region, but even at the same points, while the geographical distribution of the more marked varieties and of related species offers no conclusion as to their origin, but only as to the last great migration of the plant world, because they arose before this period, as indeed appears from their distribution.

Just as different varieties arise simultaneously from one kins.h.i.+p at the same place, the same variety may arise in places far separated, when the a.n.a.logous external exciting causes occasion an identical transformation in the idioplasm. The experimental proof lies in the fact that like beginnings of varieties often appear at great distances from each other.

An apparent social origin of varieties is indicated, when, after having come together in migration, they first develop the unlike determinants which they have gained in various locations. An apparently individual origin of the same or different varieties is indicated, when the formation of the determinants take place at one and the same place, but their development follows only after the kindred has been scattered by migration.

19. GENERAL RELATION OF THE PHYLOGENETIC LINES IN THE ORGANIC KINGDOMS.

Since the nature of an organism is contained in the sum of its idioplasmic determinants alone, the evolution of a phylogeny consists in the evolution of the idioplasm. This is perceived from the succession of the visible ontogenetic characteristics which in general run parallel with it. The idioplasm varies in two ways: (1) by an _automatic perfecting process_; (2) by _adaptation to environment_.

By virtue of the _automatic variation_ of the idioplasm the ontogenies of a phylogenetic line attain to a continually more complex organization and greater differentiation of function. In this differentiation, however, only the qualitative differences are of importance; quant.i.tative and numerical gradations may be disregarded. The more complex admits of more combinations than the simpler; hence if a phylogeny reaches a higher stage by automatic evolution it may branch into several lines, of which each appears as the continuation of the parent stock.

Since _adaptive variations_ depend only on the trans.m.u.tations of environment, an organism may rise to a higher organization and division of labor by continually adapting itself to the changed environment. But the organism may also change its adaptation while it remains at the same stage of organization. And since the adaptive variation is quickly perfected as compared with automatic evolution, although extremely slowly as compared with the duration of the ontogeny, an organization may change its adaptation several times while it remains at the same grade of organization and division of labor. Since there are also numerous different kinds of adaptation, a phyletic line may divide at each point into several adaptive forms, which appear in the taxonomic system as species, genera, often even as whole families, while in other cases various degrees of organization have appeared in one family.

20. LAWS OF EVOLUTION OF THE PLANT KINGDOM.

In the sub-organic kingdom, which precedes the plant and animal kingdoms, (see page 5), there are gradually formed from the spontaneously generated plasma independent cells with their characteristic properties, _i.e._, growth by intussusception of micellae, formation of a plasmic cuticle, and a non-plasmic membrane about the same, division of the cells, separation of the cells thus formed, and free cell formation within the cell contents. These properties are inherited from the sub-organic kingdom by the plants and animals which follow in the next stage of phylogeny. The evolution of the plant kingdom proceeds through the following regular processes, which continue to operate through the entire phylogenetic series.

_Law of Phylogenetic Combination._--The simplest of all plants are cells of round form, which grow and reproduce themselves by division, budding or free cell formation. From the fact that the younger generation of cells, instead of separating from each other and growing to independent plant individuals, remain united with each other, multicellular plants arise from unicellular. The same transformation of the reproductive cells into non-separable tissue cells is repeated several times in multicellular plants and serves to enlarge the individual. There is manifested in this phylogenetic process the tendency of the plant to combine in the higher stages into one complex whole those parts which in the lower stages tend to be independent. A similar unifying tendency is revealed also in those plant members which have arisen by differentiation and represent a system only by their being connected at certain points. These combine in the higher stages and form ultimately continuous tissues.

_Law of Phylogenetic Complication or Ampliation, Differentiation and Reduction._--The cells, and, in general, the parts of plants which lie near each other in s.p.a.ce or follow upon each other in time, are always alike in the lower stages. By differentiation they become unlike, so that the sum of the functions which at first fall to the lot of all parts without distinction now is shared among the individual parts. By this means each part can perform its own special function so much the better. Differentiation is repeated in the course of the phylogeny, since at first all parts of an ontogeny diverge into two or more parts, then the parts of these parts divide again, etc. Along with this process of division another process is always active, which, as it were, prepares the way for the former, namely, ampliation, by virtue of which the growth of the whole ontogeny or of single stages of it undergoes a quant.i.tative increase, so that an organ acquires a greater number of cells, and an individual a greater number of organs. After this increase in number of parts in a stage of ontogeny, differentiation follows as far as the nature of the functions permits, by the parts most separated pa.s.sing into each other by intermediate gradations. By the further phylogenetic process of reduction the intermediate forms are suppressed.

At last only the extreme products of differentiation lie near each other in s.p.a.ce or follow upon each other in time; and these products are as limited in quant.i.ty and number as possible.

Along with the above named phylogenetic processes, which take place by the automatic increase of the idioplasm, external influences are always active. These lend to the organism at times a local stamp corresponding to its environment, and follow the law of adaptation.

21. ALTERNATION OF GENERATIONS IN RELATION TO PHYLOGENY.

Since the simplest plants are cells and the more complex ones are formed from cells, a whole phylogenetic line may be regarded as a series of cell generations following one after another. In the lowest forms all cell generations are like each other; in all others they show differences which become continually greater and more numerous. Thus alternation of generations in cells exists, because the successive generations become more and more complicated at each succeeding period.

Among these periods the ontogenetic period or ontogeny embraces all generations from one cell to the return of the exactly similar kind of cell. In the lowest forms of cell differentiation the cells of successive generations are all independent; the ontogenetic period consists of a cycle of generations of unicellular plants. Later the cell generations of an ontogeny are united by parts into plant individuals; the ontogenetic period consists of a cycle of multicellular and unicellular, or only of multicellular plant generations. If all the cell generations of an ontogenetic period have been united into a single individual, the successive plant generations are alike and alternation of generations has ceased.

The unlikeness of the generations arises either from inner causes of temporary differentiation alone, or by temporary differentiation which receives a definite imprint by the change of seasons. But in the latter case the characteristic of adaptation is again lost in the course of the phylogeny and alternation of generations follows then without regard to the season. If the given adaptation is united in the lower plants with alternation of generations during the ontogenetic periods, one of the unlike plant generations is repeated an indefinite number of times (repet.i.tional generation), while the other unlike plant generation appears only once and then at the beginning of the resting stage and remains latent in the form of a resting spore till the beginning of the next period of generation. With this peculiar transition generation, which has arisen in the lower stages as.e.xually, and in the following higher stages by the union of a male and a female cell, and which hence is hermaphrodite, there are generally a.s.sociated later two other single generations--_viz._, a generation preceding and one following the hermaphrodite, the former as a s.e.x-producing generation, the other as a s.e.x-produced generation.

The phylogenetic significance of the alternation of generations consists in its representing a transition stage from the unicellular to the simpler multicellular and from the latter to the more complex multicellular plants. The plant generations of any phylogenetic stage increase by ampliation, become unlike by differentiation in time (alternation of generations), and unite in a plant individual, whose unlike ontogenetic stages correspond to the unlike plant generations of the earlier ancestral series.

22. MORPHOLOGY AS THE SCIENCE OF PHYLOGENY.

All organic phenomena belong, according to their causes, to two different cla.s.ses: (1) Those belonging to one group are the results of external influences in each ontogeny and are not inherited; they represent nutrition varieties, are experimentally demonstrable, and const.i.tute the subject matter of experimental physiology. (2) The others are inherited and again transmitted; they belong to the physiology of the idioplasm. This subject is mainly occupied with the origin of the determinants, hence with the formation of varieties and species. It is not the subject of experiment, and const.i.tutes the phylogeny or the physiology of the formation of determinants. A sub-division of this subject is occupied with the development of the determinants already present, hence with the formation of races. It is elucidated especially by experiments in crossing and may be designated as the physiology of the development of the determinants.

The morphological phenomena which find their application in taxonomy, belong exclusively to phylogeny. Their ontogenetic history does not explain their true significance; this can be known only in a phylogenetic way by comparison of one phenomenon with those phenomena from which it has arisen in the course of evolution.

23. PLANT CLa.s.sIFICATION FROM THE STANDPOINT OF PHYLOGENY.

Spontaneous generation has taken place at all times and in all places, in as far as the necessary conditions were concurrently present. (See page 47). After spontaneous generation the automatic phylogenetic evolution begins and advances constantly. Consequently the phylogenetic line rises from time to time to higher stages of organization and division of labor, but dies of old age if the automatic perfecting process ceases. The phylogenetic lines of organisms now living have therefore an unequal age; those of the most highly developed plants and animals had their origin in the earliest periods of organic life, those of the lowest organisms in the most recent periods. Hence no general genetic relation exists among lines now living; only those that are nearly related and have reached approximately equal stages of organization may be regarded as branches of the same phylogenetic stock.

A phylogenetic plant system does not exist in fact, but only in figure.

If genetic relation between two races is a.s.sumed, either as a reality or as a symbol, the degree of relations.h.i.+p is determined in a theoretically exact manner by the number and length of the phylogenetic steps which are found either between them both or between them and the common starting point, according as races belong to the same or collateral lines. The fact that two organisms belong to the same line of descent is recognized from the ontogeny of the higher including the ontogeny of the lower.

Since only a proportionately small number of known forms can appear as types of the supposed stages of evolution, only a few phylogenetic lines, and these only in a general way, may be established, on account of the great incompleteness of the present plant world. Such a line proceeds from the green filamentous algae through the liverworts to the vascular plants. Among the phanerogams, apparently so numerously represented, only phylogenetic series of individual organs can be ascertained, but no phylogenetic series of families. A phylogenetic system of phanerogams is not to be hazarded in the roughest outline.

Even the relative rank of the two chief divisions of the angiosperms, the monocotyledons and dicotyledons, is a matter of question, as also which family in each of these divisions is to be considered the most perfect.