The Principles of Biology Vol. I/Chapter II.6
CHAPTER VI.
INDIVIDUALITY.
§ 72. What is an individual? is a question which many readers will think it easy to answer. Yet it is a question that has led to much controversy among Zoologists and Botanists, and no quite satisfactory reply to it seems possible. As applied to a man, or to any one of the higher animals, which are all sharply-defined and independent, the word individual has a clear meaning: though even here, when we turn from average cases to exceptional cases—as a calf with two heads and two pairs of fore-limbs—we find ourselves in doubt whether to predicate one individuality or two. But when we extend our range of observation to the organic world at large, we find that difficulties allied to this exceptional one meets us everywhere under every variety of form.
Each uniaxial plant may perhaps fairly be regarded as a distinct individual; though there are botanists who do not make even this admission. What, however, are we to say of a multiaxial plant? It is, indeed, usual to speak of a tree with its many branches and shoots as singular; but strong reasons may be urged for considering it as plural. Every one of its axes has a more or less independent life, and when cut off and planted may grow into the likeness of its parent; or, by grafting and budding, parts of this tree may be developed upon another tree, and there manifest their specific peculiarities. Shall we regard all the growing axes thus resulting from slips and grafts and buds, as parts of one individual or as distinct individuals? If a strawberry-plant sends out runners carrying buds at their ends, which strike root and grow into independent plants that separate from the original one by decay of the runners, must we not say that they possess separate individualities; and yet if we do this, are we not at a loss to say when their separate individualities were established, unless we admit that each bud was from the beginning an individual? Commenting on such perplexities Schleiden says—"Much has been written and disputed concerning the conception of the individual, without, however, elucidating the subject, principally owing to the misconception that still exists as to the origin of the conception. Now the individual is no conception, but the mere subjective comprehension of an actual object, presented to us under some given specific conception, and on this latter it alone depends whether the object is or is not an individual. Under the specific conception of the solar system, ours is an individual: in relation to the specific conception of a planetary body, it is an aggregate of many individuals." ... "I think, however, that looking at the indubitable facts already mentioned, and the relations treated of in the course of these considerations, it will appear most advantageous and most useful, in a scientific point of view, to consider the vegetable cell as the general type of the plant (simple plant of the first order). Under this conception, Protococcus and other plants consisting of only one cell, and the spore and pollen-granule, will appear as individuals. Such individuals may, however, again, with a partial renunciation of their individual independence, combine under definite laws into definite forms (somewhat as the individual animals do in the globe of the Volvox globator[1]). These again appear empirically as individual beings, under a conception of a species (simple plants of the second order) derived from the form of the normal connexion of the elementary individuals. But we cannot stop here, since Nature herself combines these individuals, under a definite form, into larger associations, whence we draw the third conception of the plant, from a connexion, as it were, of the second power (compound plants—plants of the third order). The simple plant proceeding from the combination of the elementary individuals is then termed a bud (gemma), in the composition of plants of the third order."
The animal kingdom presents still greater difficulties. When, from sundry points on the body of a common polype, there bud out young polypes which, after acquiring mouths and tentacles and closing up the communications between their stomachs and the stomach of the parent, finally separate from the parent; we may with propriety regard them as distinct individuals. But when in the allied compound Hydrozoa, we find that these young polypes continue permanently connected with the parent; and when by this continuous budding-out there is presently produced a tree-like aggregation, having a common alimentary canal into which the digestive cavity of each polype opens; it is no longer so clear that these little sacs, furnished with mouths and tentacles, are severally to be regarded as distinct individuals. We cannot deny a certain individuality to the polypedom. And on discovering that some of the buds, instead of unfolding in the same manner as the rest, are transformed into capsules in which eggs are developed—on discovering that certain of the incipient polypes thus become wholly dependent on the aggregate for their nutrition, and discharge functions which have nothing to do with their own maintenance, we have still clearer proof that the individualities of the members are partially merged in the individuality of the group. Other organisms belonging to the same order, display still more decidedly this transition from simple individualities to a complex individuality. In the Diphyes there is a special modification of one or more members of the polypedom into a swimming apparatus which, by its rhythmical contractions, propels itself through the water, drawing the polypedom after it. And in the more differentiated Physalia various organs result from the metamorphosis of parts which are the homologues of individual polypes. In this last instance, the individuality of the aggregate is so predominant that the individualities of its members are practically lost. This combination of individualities in such way as to produce a composite individual, meets us in other forms among the ascidians. While in some of these, as in the Clavelina and in the Botryllidæ, the animals associated are but little subordinated to the community they form, in others they are so combined as to form a compound individual. The pelagic ascidian Doliolum is an example. "Here we find a large individual which swims by contractions of circular muscular bands, carries a train of smaller individuals attached to a long dorsal process of the test. These are arranged in three rows: those constituting the lateral row have wide mouths and no sexual organs or organs of locomotion—they subserve the nutrition of the colony, a truth which is illustrated by the fact that as soon as they are properly developed the large individual (the mother) loses her alimentary canal;" while from the median row are eventually derived the sexual zoids.
On the hypothesis of Evolution, perplexities of this nature are just such as we might anticipate. If Life in general commenced with minute and simple forms, like those out of which all organisms, however complex, now originate; and if the transitions from these primordial units to organisms made up of groups of such units, and to higher organisms made up of groups of such groups took place by degrees; it is clear that individualities of the first and simplest order would merge gradually in those of a larger and more complex order, and these again in others of an order having still greater bulk and organization. Hence it would be impossible to say where the lower individualities ceased and the higher individualities commenced.
§ 73. To meet these difficulties, it has been proposed that the whole product of a single fertilized germ shall be regarded as a single individual; whether such whole product be organized into one mass, or whether it be organized into many masses that are partially or completely separate. It is urged that whether the development of the fertilized germ be continuous or discontinuous ( § 50) is a matter of secondary importance; that the totality of living tissue to which the fertilized germ gives rise in any one case, is the equivalent of the totality to which it gives rise in any other case; and that we must recognize this equivalence, whether such totality of living tissue takes a concrete or a discrete arrangement. In pursuance of this view, a zoological individual is constituted either by any such single animal as a mammal or bird, which may properly claim the title of a zoon, or by any such group of animals as the numerous Medusæ that have been developed from the same egg, which are to be severally distinguished as zooids.
Admitting it to be very desirable that there should be words for expressing these relations and this equivalence, it may be objected that to apply the word individual to a number of separate living bodies, is inconvenient: conflicting so much, as it does, with the ordinary conception which this word suggests. It seems a questionable use of language to say that the countless masses of Anacharis Alsinastrum (now Eloidea canadensis) which, within these few years, have grown up in our rivers, canals, and ponds, are all parts of one individual: and yet as this plant does not seed in England, these countless masses, having arisen by discontinuous development, must be so regarded if we accept the above definition.
It may be contended, too, that while it does violence to our established way of thinking, this mode of interpreting the facts is not without its difficulties. Something seems to be gained by restricting the application of the title individual, to organisms which, being in all respects fully developed, possess the power of producing their kind after the ordinary sexual method, and denying this title to those incomplete organisms which have not this power. But the definition does not really establish this distinction for us. On the one hand, we have cases in which, as in the working bee, the whole of the germ-product is aggregated into a single organism; and yet, though an individual according to the definition, this organism has no power of reproducing its kind. On the other hand, we have cases like that of the perfect Aphis, where the organism is but an infinitesimal part of the germ product, and yet has that completeness required for sexual reproduction. Further, it might be urged with some show of reason, that if the conception of individuality involves the conception of completeness, then, an organism which possesses an independent power of reproducing itself, being more complete than an organism in which this power is dependent on the aid of another organism, is more individual.
§ 74. There is, indeed, as already implied, no definition of individuality that is unobjectionable. All we can do is to make the best practicable compromise.
As applied either to an animate or an inanimate object, the word individual ordinarily connotes union among the parts of the object and separateness from other objects. This fundamental element in the conception of individuality, we cannot with propriety ignore in the biological application of the word. That which we call an individual plant or animal must, therefore, be some concrete whole and not a discrete whole. If, however, we say that each concrete living whole is to be regarded as an individual, we are still met by the question—What constitutes a concrete living whole? A young organism arising by internal or external gemmation from a parent organism, passes gradually from a state in which it is an indistinguishable part of the parent organism to a state in which it is a separate organism of like structure with the parent. At what stage does it become an individual? And if its individuality be conceded only when it completely separates from the parent, must we deny individuality to all organisms thus produced which permanently retain their connexions with their parents? Or again, what must we say of the Hectocotylus, which is an arm of the Cuttle-fish that undergoes a special development and then, detaching itself, lives independently for a considerable period? And what must we say of the larval nemertine worm the pilidium of which with its nervous system is left to move about awhile after the developing worm has dropped out of it?
To answer such questions we must revert to the definition of life. The distinction between individual in its biological sense, and individual in its more general sense, must consist in the manifestation of Life, properly so called. Life we have seen to be, "the definite combination of heterogeneous change, both simultaneous and successive, in correspondence with external co-existences and sequences." Hence, a biological individual is any concrete whole having a structure which enables it, when placed in appropriate conditions, to continuously adjust its internal relations to external relations, so as to maintain the equilibrium of its functions. In pursuance of this conception, we must consider as individuals all those wholly or partially independent organized masses which arise by multicentral and multiaxial development that is either continuous or discontinuous ( § 50). We must accord the title to each separate aphis, each polype of a polypedom, each bud or shoot of a lowering plant, whether it detaches itself as a bulbil or remains attached as a branch.
By thus interpreting the facts we do not, indeed, avoid all anomalies. While, among flowering plants, the power of independent growth and development is usually possessed only by shoots or axes; yet, in some cases, as in that of the Begonia-leaf awhile since mentioned, the appendage of an axis, or even a small fragment of such appendage, is capable of initiating and carrying on the functions of life; and in other cases, as shown by M. Naudin in the Drosera intermedia, young plants are occasionally developed from the surfaces of leaves. Nor among forms like the compound Hydrozoa, does the definition enable us to decide where the line is to be drawn between the individuality of the group and the individualities of the members: merging into each other, as these do, in different degrees. But, as before said, such difficulties must necessarily present themselves if organic forms have arisen by insensible gradations. We must be content with a course which commits us to the smallest number of incongruities; and this course is, to consider as an individual any organized mass which is capable of independently carrying on that continuous adjustment of inner to outer relations which constitutes Life.
- ↑ Whether the Volvox is to be classed as animal or vegetal is a matter of dispute; but its similarity to the blastula stage of many animals warrants the claim of the zoologists.