Popular Science Monthly/Volume 29/June 1886/The Factors of Organic Evolution III
|THE FACTORS OF ORGANIC EVOLUTION.|
LIMITED, as thus far drawn, to a certain common trait of those minute organisms which are mostly below the reach of unaided vision, the foregoing conclusion appears trivial enough. But it ceases to appear trivial on passing beyond these limits, and observing the implications, direct and indirect, as they concern plants and animals of sensible sizes.
Popular expositions of science have so far familiarized many readers with a certain fundamental trait of living things around, that they have ceased to perceive how marvellous a trait it is, and until interpreted by the Theory of Evolution, how utterly mysterious. In past times, the conception of an ordinary plant or animal which prevailed, not throughout the world at large only but among the most instructed, was that it is a single continuous entity. One of these living things was unhesitatingly regarded as being in all respects a unit. Parts it might have, various in their sizes, forms, and compositions; but these were components of a whole which had been from the beginning in its original nature a whole. Even to naturalists fifty years ago, the assertion that a cabbage or a cow, though in one sense a whole, is in another sense a vast society of minute individuals, severally living in greater or less degrees, and some of them maintaining their independent lives unrestrained, would have seemed an absurdity. But this truth which, like so many of the truths established by science, is contrary to that common sense in which most people have so much confidence, has been gradually growing clear since the days when Leeuwenhoeck and his contemporaries began to examine through lenses the minute structures of common plants and animals. Each improvement in the microscope, while it has widened our knowledge of those minute forms of life described above, has revealed further evidence of the fact that all the larger forms of life consist of units severally allied in their fundamental traits to these minute forms of life. Though, as formulated by Schwann and Schleiden, the cell-doctrine has undergone qualifications of statement; yet the qualifications have not been such as to militate against the general proposition that organisms visible to the naked eye, are severally compounded of invisible organisms—using that word in its most comprehensive sense. And then, when the development of any animal is traced, it is found that having been primarily a nucleated cell, and having afterwards become by spontaneous fission a cluster of nucleated cells, it goes on through successive stages to form out of such cells, ever multiplying and modifying in various ways, the several tissues and organs composing the adult.
On the hypothesis of evolution this universal trait has to be accepted not as a fact that is strange but unmeaning. It has to be accepted as evidence that all the visible forms of life have arisen by union of the invisible forms; which, instead of flying apart, when they divided, remained together. Various intermediate stages are known. Among plants, those of the Volvox type show us the component protophytes so feebly combined that they severally carry on their lives with no appreciable subordination to the life of the group. And among animals, a parallel relation between the lives of the units and the life of the group is shown us in Uroglena and Syncrypta. From these first stages upwards, may be traced through successively higher types, an increasing subordination of the units to the aggregate; though still a subordination leaving to them conspicuous amounts of individual activity. Joining which facts with the phenomena presented by the cell-multiplication and aggregation of every unfolding germ, naturalists are now accepting the conclusion that by this process of composition from Protozoa were formed all classes of the Metazoa—(as animals formed by this compounding are now called); and that in a similar way from Protophyta, were formed all classes of what, by analogy, I suppose will be called Metaphyta, though the word does not yet seem to have become current.
And now what is the general meaning of these truths, taken in connexion with the conclusion reached in the last section? It is that this universal trait of the Metazoa and Metaphyta, must be ascribed to the primitive action and re-action between the organism and its medium. The operation of those forces which produced the primary differentiation of outer from inner in early minute masses of protoplasm, pre-determined this universal cell-structure of all embryos, plant and animal, and the consequent cell-composition of adult forms arising from them. How unavoidable is this implication, will be seen on carrying further an illustration already—used that of the shingle covered shore, the pebbles on which, while being in some cases selected, have been in all cases rounded and smoothed. Suppose a bed of such shingle to be, as we often see it, solidified, along with interfused material, into a conglomerate. What in such case must be considered as the chief trait of such conglomerate; or rather—what must we regard as the chief cause of its distinctive characters? Evidently the action of the sea. Without the breakers, no pebbles; without the pebbles, no conglomerate. Similarly then, in the absence of that action of the medium by which was effected the differentiation of
outer from inner in those microscopic portions of protoplasm constituting the earliest and simplest animals and plants, there could not have existed this cardinal trait of composition which all the higher animals and plants show us.
So that, active as has been the part played by natural selection, alike in modifying and moulding the original units—largely as survival of the fittest has been instrumental in farthering and controlling the a cerebration of these units into visible organisms and eventually into large ones: ye: we must ascribe to the direct effect of the medium on primitive forms of life, that primordial trait of which this everywhere-operative factor has taken advantage.
Let us turn now: a another and more manifest trait of higher organisms, for which also there is this same general cause. Let us observe how, on a higher platform, there recurs this differentiation of outer from inner how this primary trait in the living units with which life commences, re-appears as a primary trait in those aggregates of units which constitute visible organisms.
In its simplest and most unmistakable form, we see this in the early changes of an unfolding ovum of primitive type. The original fertilized single cell, having by spontaneous fusion multiplied into a cluster of such cells, there begins to show itself a contrast between periphery and centre; and presently there is formed a sphere consisting of a superficial layer unlike its contents. The first change, then, is the rise of a difference between that outer part which holds direct converse with the surrounding medium, and that inclosed part which does not. This primary differentiation in these compound embryos of higher animals, parallels the primary differentiation undergone by the simplest living things.
Leaving, for the present, succeeding changes of the compound embryo, the significance of which we shall have to consider by-and-by. let us pass now to the adult forms of visible plants and animals. In them we find cardinal traits which, after what we have seen above, will further impress us with the importance of the effects wrought on the organism by its medium.
From the thallus of a sea-weed up to the leaf of a highly developed phænogam, we find, at all stages, a contrast between the inner and outer parts of these flattened masses of tissue. In the higher Algæ "the outermost layers consist of smaller and firmer cells, while the inner cells are often very large, and sometimes extremely long;" and in the leaves of trees the epidermal layer, besides differing in the sizes and shapes of its component cells from the parenchyma forming the inner substance of the leaf, is itself differentiated by having a continuous continuous cuticle, and by Laving the outer walls of its cells unlike the inner walls. Especially instructive is the structure of such intermediate types as the Liverworts. Beyond the differentiation of the raring cells from the contained cells, and the contrast between upper surface and under surface, the frond of Marchantia polymorpha clearly shows us the direct effect of incident forces: and shows us, too, how it is involved with the effect of inherited proclivities. The frond grows from a flat disc-shaped gemma, the two sides of which are alike. Either side may fall uppermost; and then of the developing shoot, the side exposed to the light "is under all circumstances the upper side which forms stomata, the dark side becomes the under side which produces root-hairs and leafy processes." So that while we have undeniable proof that the contrasted influences of the medium on the two sides, initiate the differentiation, we have also proof that the completion of it is determined by the transmitted structure of the type; since it is impossible to ascribe the development of stomata to the direct action of air and light. On turning from foliar expansion to stems and roots, facts of like meaning meet us. Speaking generally of epidermal tissue and inner tissue, Sachs remarks that "the contrast of the two is the plainer the more the part of the plant concerned is exposed to air and light." Elsewhere, in correspondence with this, it is said that in roots the cells of the epidermis, though distinguished by bearing hairs, "are otherwise similar to those of the fundamental tissue" which they clothe, while the cuticular covering is relatively thin; whereas in stems the epidermis (often further differentiated) is composed of layers of cells which are smaller and thicker-walled: a stronger contrast of structure corresponding to a stronger contrast of conditions. By way of meeting the suggestion that these respective differences are wholly due to the natural selection of favourable variations, it will suffice if I draw attention to the unlikeness between imbedded roots and exposed roots. While in darkness, and surrounded by moist earth, the outermost protective coats, even of large roots, are comparatively thin; but when the accidents of growth entail permanent exposure to light and air, roots acquire coverings allied in character to the coverings of branches. That the action of the medium causes these and converse changes, cannot be doubted when we find, on the one hand, that "roots can become directly transformed into leaf-bearing shoots," and, on the other hand, that in some plants certain "apparent roots are only underground shoots," and that nevertheless "they are similar to true roots in function and in the formation of tissue, but have no root-cap, and, when they come to the light above ground, continue to grow in the manner of ordinary leaf-shoots." If, then, in highly developed plants inheriting pronounced structures, this differentiating influence of the medium is so marked, it must have been all-important at the outset while types were undetermined.
As with plants so with animals, we find good reason for inferring that while all the specialities of the tegumentary parts must be ascribed to the natural selection of favourable variations, their most general traits are due to the direct action of surrounding agencies. Here we come upon the border of those changes which are ascribable to use and disuse. But from this class of changes we may fitly exclude those in which the parts concerned are wholly or mainly passive. A corn and a blister will conveniently serve to illustrate the way in which certain outer actions produce in the superficial tissues, effects of a purely physical kind—effects related neither to the needs of the organism nor to its structural proclivities. They are neither adaptive changes nor changes towards completion of the type. After noting them we may pass to allied, but still more instructive, facts. Continuous pressure on any portion of the surface causes absorption, while intermittent pressure causes growth: the one impeding circulation and the passage of plasma from the capillaries into the tissues, and the other aiding both. There are further mechanically-produced effects. That the general character of the ribbed skin on the under surfaces of the feet and insides of the hands is directly due to friction and intermittent pressure, we have the proofs:—first, that the tracts most exposed to rough usage are the most ribbed; second, that the insides of hands subject to unusual amounts of rough usage, as those of sailors, are strongly ribbed all over; and third, that in hands which are very little used, the parts commonly ribbed become quite smooth. These several kinds of evidence, however, full of meaning as they are, I give simply to prepare the way for evidence of a much more conclusive kind.
Where ulceration has eaten away the deep-seated layer out of which the epidermis grows, or where this layer has been destroyed by an extensive burn, the process of healing is very significant. From the subjacent tissues, which in the normal order have no concern with outward growth, there is produced a new skin, or rather a pro-skin; for this substituted outward-growing layer contains no hair-follicles or other specialities of the original one. Nevertheless, it is like the original one in so far that it is a continually renewed protective covering. Doubtless it may be contended that this make-shift skin results from the inherited proclivity of the type—the tendency to complete afresh the structure of the species when injured. We cannot, however, ignore the immediate influence of the medium, on recalling the facts above named, or on remembering the further fact that an inflamed surface of skin, when not sheltered from the air, will throw out a film of coagulable lymph. But that the direct action of the medium is a chief factor we are clearly shown by another case. Accident or disease occasionally causes permanent eversion, or protrusion, of mucous membrane. After a period of irritability, great at first but decreasing as the change advances, this membrane assumes the general character of ordinary skin. Nor is this all: its microscopic structure changes. Where it is a mucous membrane of the kind covered by cylinder-epithelium, the cylinders gradually shorten, becoming finally flat, and there results a squamous epithelium: there is a near approach in minute composition to epidermis. Here a tendency towards completion of the type cannot be alleged; for there is, contrariwise, divergence from the type. The effect of the medium is so great that, in a short time, it overcomes the inherited proclivity and produces a structure of opposite kind to the normal one.
Fully to perceive the way in which these evidences compel us to recognize the influence of the medium as a primordial factor, we need but conceive them as interpreted without it. Suppose, for instance, we say that the structure of the epidermis is wholly determined by the natural selection of favourable variations; what must be the position taken in presence of the fact above named, that the cell-structure of mucous membrane changes into the cell-structure of skin when mucous membrane is exposed to the air? The position taken must be this:—Though mucous membrane in a highly-evolved individual organism, thus shows the powerful effect of the medium on its surface; yet we must not suppose that the medium had the effect of producing such a cell-structure on the surfaces of primitive forms, undifferentiated though they were; or, if we suppose that such an effect was produced on them, we must not suppose that it was inheritable. Contrariwise, we must suppose that such effects of the medium either were not wrought at all, or that they were evanescent: though repeated through millions upon millions of generations they left no traces. And we must conclude that this skin-structure arose only in consequence of spontaneous variations not physically initiated (though like those physically initiated) which natural selection laid hold of and increased. Does any one think this a tenable position?
And now we approach the last and chief series of morphological phenomena which must be ascribed to the direct action of environing matters and forces. These are presented to us when we study the early stages in the development of the embryos of the Metazoa in general.
We will set out with the fact already noted in passing, that after repeated spontaneous fissions have changed the original fertilized germ cell into that cluster of cells which forms a gemmule or a primitive ovum, the first contrast which arises is between the peripheral parts and the central parts. Where, as with lower creatures which do not lay up large stores of nutriment with the germs of their offspring, the inner mass is inconsiderable, the outer layer of cells, which are presently made quite small by repeated subdivisions, forms a membrane extending over the whole surface—the blastoderm. The next stage of development, which ends in this covering layer becoming double, is reached in two ways—by invagination and by delamination; but which is the original way, and which the abridged way, is not quite certain. Of invagination, multitudinously exemplified in the lowest types, Mr. Balfour says:—"On purely à priori grounds there is in my opinion more to be said for invagination than for any other view"; and, for present purposes, it will suffice if we limit ourselves to this: making its nature clear to the general reader by a simple illustration.
Take a small india-rubber ball—not of the inflated kind, nor of the solid kind, but of the kind about an inch or so in diameter with a small hole through which, under pressure, the air escapes. Suppose that instead of consisting of india-rubber its wall consists of small cells made polyhedral in form by mutual pressure, and united together. This will represent the blastoderm. Now with the finger, thrust in one side of the ball until it touches the other: so making a cup. This action will stand for the process of invagination. Imagine that by continuance of it, the hemispherical cup becomes very much deepened and the opening narrowed, until the cup becomes a sac, of which the introverted wall is everywhere in contact with the outer wall. This will represent the two-layered "gastrula"—the simplest ancestral form of the Metazoa: a form which is permanently represented in some of the lowest types; for it needs but tentacles round the mouth of the sac, to produce a common hydra. Here the fact which it chiefly concerns us to remark, is that of these two layers the outer, called in embryological language the epiblast, continues to carry on direct converse with the forces and matters in the environment; while the inner, called the hypoblast, comes in contact with such only of these matters as are put into the food-cavity which it lines. We have further to note that in the embryos of Metazoa at all advanced in organization, there arises between these two layers a third—the mesoblast. The origin of this is seen in types where the developmental process is not obscured by the presence of a large food-yolk. While the above-described introversion is taking place, and before the inner surfaces of the resulting epiblast and hypoblast have come into contact, cells, or amœboid units equivalent to them, are budded off from one or both of these inner surfaces, or some part of one or other; and these form a layer which eventually lies between the other two—a layer which, as this mode of formation implies, never has any converse with the surrounding medium and its contents, or with the nutritive bodies taken in from it. The striking facts to which this description is a necessary introduction, may now be stated. From the outer layer, or epiblast, are developed the permanent skin and its out-growths, the nervous system, and the organs of sense; from the introverted layer, or hypoblast, are developed the alimentary canal along with those parts of its appended organs, liver, pancreas, &c., which are concerned in delivering their secretions into the alimentary canal, as well as the linings of those ramifying tubes in the lungs which convey air to the places where gaseous exchange is effected. And from the mesoblast originate the bones, the muscles, the heart and blood-vessels, and the lymphatics, together with such parts of various internal organs as are most remotely concerned with the outer world. Minor qualifications being admitted, there remain the broad general facts, that out of that part of the external layer which remains permanently external, are developed all the structures which carry on intercourse with the medium and its contents, active and passive; out of the introverted part of this external layer, are developed the structures which carry on intercourse with the quasi-external substances that are taken into the interior—solid food, water, and air; while out of the mesoblast are developed structures which have never had, from first to last, any intercourse with the environment. Let us contemplate these general facts.
Who would have imagined that the nervous system is a modified portion of the primitive epidermis? In the absence of proofs furnished by the concurrent testimony of embryologists during the last thirty or forty years, who would have believed that the brain arises from an infolded tract of the outer skin, which, sinking down beneath the surface, becomes imbedded in other tissues and eventually surrounded by a bony case? Yet the human nervous system in common with the nervous systems of lower animals is thus originated. In the words of Mr. Balfour, early embryological changes imply that—
Less startling perhaps, though still startling enough, is the fact that the eye is evolved out of a portion of the skin; and that while the crystalline lens and its surroundings thus originate, the "percipient portions of the organs of special sense, especially of optic organs, are often formed from the same part of the primitive epidermis" which forms the central nervous system. Similarly is it with the organs for smelling and hearing. These, too, begin as sacs formed by infoldings of the epidermis; and while their parts are developing they are joined from within by nervous structures which were themselves epidermic in origin. How are we to interpret these strange transformations? Observing, as we pass, how absurd from the point of view of the special-creationist, would appear such a filiation of structures, and such a round-about mode of embryonic development, we have here to remark that the process is not one to have been anticipated as a result of natural selection. After numbers of spontaneous variations had occurred, as the hypothesis implies, in useless ways, the variation which primarily initiated a nervous centre might reasonably have been expected to occur in some internal part where it would be fitly located: its initiation in a dangerous place and subsequent migration to a safe place, would be incomprehensible. Not so if we bear in mind the cardinal truth above set forth, that the structures for holding converse with the medium and its contents, arise in that completely superficial part which is directly affected by the medium and its contents; and if we draw the inference that the external actions themselves initiate the structures. These once commenced, and furthered by natural selection where favourable to life, would form the first term of a series ending in developed sense organs and a developed nervous system.
Though it would enforce the argument, I must, for brevity's sake, pass over the analogous evolution of that introverted layer, or hypoblast, out of which the alimentary canal and attached organs arise. It will suffice to emphasize the fact that having been originally external, this layer continues in its developed form to have a quasi-externality, alike in its digesting part and in its respiratory part; since it continues to deal with matters alien to the organism. I must also refrain from dwelling at length on the fact already adverted to, that the intermediate derived layer, or mesoblast, which was at the outset completely internal, originates those structures which ever remain completely internal, and have no communication with the environment save through the structures developed from the other two: an antithesis which has great significance.
Here, instead of dwelling on these details, it will be better to draw attention to the most general aspect of the facts. Whatever may be the course of subsequent changes, the first change is the formation of a superficial layer or blastoderm; and by whatever series of transformations the adult structure is reached, it is from the blastoderm that all the organs forming the adult originate. Why this marvellous fact? Why out of the primitive mass of organizable substance which is to form a new creature, should its surface be the part from which is remotely derived its entire structure? Before embryologists had established this truth, anyone who had asserted it would have been thought insane; and even now it remains a mystery if we refuse to take account of the direct relations between the organism and the medium. But we need only consider the incidents of this relation to get a feasible explanation. Before yet the primitive metazoon had any structure beyond that possessed by its component cells, its outer surface was the part through which nutritive matters were taken in and through which were absorbed and exhaled, oxygen and carbonic acid. Its outer surface was the part which now touched quiescent masses, and now received the collisions consequent on its own motions or the motions of others similarly carried along by their cilia. Its outer surface was the part to receive the sound-vibrations occasionally propagated through the water; the part to be affected more strongly than any other by those variations in the amounts of light caused by the passing of small bodies close to it; and the part which met those diffused molecules constituting odours. That is to say, at the outset the surface was the part on which there fell the various influences pervading the environment, through which there passed the materials for growth furnished by the environment, by which there were received those impressions from the environment serving for the guidance of actions, and which had to bear the mechanical re-actions consequent upon such actions. Necessarily, therefore, the surface was the part in which were initiated the various instrumentalities for carrying on intercourse with the environment. To suppose otherwise is to suppose that such instrumentalities arose internally where they could neither be operated on by surrounding agencies nor operate on them,—where the differentiating forces did not come into play, and the differentiated structures had nothing to do; and it is to suppose that meanwhile the parts directly exposed to the differentiating forces remained unchanged. Clearly, then, organization could not but begin on the surface; and having thus begun, its subsequent course could not but be determined by its superficial origin. And hence these remarkable facts showing us that individual evolution is accomplished by successive in-foldings and in-growings. Doubtless natural selection soon came into action, as, for example, in the removal of the rudimentary nervous centres from the surface; since an individual in which they were a little more deeply seated would be less likely to be incapacitated by injury of them. And so in multitudinous other ways. But nevertheless, as we here see, natural selection could operate only under subjection: it could do no more than take advantage of those structural changes which the medium and its contents initiated.
See, then, how large has been the part played by this primordial factor. Had it done no more than give to Protozoa and Protophyta that cell-form which characterizes them—had it done no more than entail the cellular composition which is so remarkable a trait of Metazoa and Metaphyta—had it done no more than cause the repetition in all visible animals and plants of that primary differentiation of outer from inner which it first wrought in animals and plants invisible to the naked eye; it would have done much towards giving to organisms of all kinds certain leading traits. But it has done more than this. By causing the first differentiations of those clusters of units out of which visible animals in general arose, it fixed the starting place for organization, and therefore determined the course of organization; and, doing this, gave indelible traits to embryonic transformations and to adult structures.
Though mainly carried on after the inductive method, the argument at the close of the foregoing section has verged towards the deductive. Here let us follow for a space the deductive method pure and simple. Doubtless in biology à priori reasoning is dangerous; but there can be no danger in considering whether its results coincide with those reached by reasoning à posteriori.
Biologists in general agree that in the present state of the world, no such thing happens as the rise of a living creature out of non-living matter. They do not deny, however, that at a remote period in the past, when the temperature of the Earth's surface was much higher than at present, and other physical conditions were unlike those we know, inorganic matter, through successive complications, gave origin to organic matter. So many substances once supposed to belong exclusively to living bodies, have now been formed artificially, that men of science scarcely question the conclusion that there are conditions under which, by yet another step of composition, quaternary compounds of lower types pass into those of highest types. That there once took place gradual divergence of the organic from the inorganic, is, indeed, a necessary implication of the hypothesis of Evolution, taken as a whole; and if we accept it as a whole, we must put to ourselves the question—What were the early stages of progress which followed, after the most complex form of matter had arisen out of forms of matter a degree less complex?
At first, protoplasm could have had no proclivities to one or other arrangement of parts; unless, indeed, a purely mechanical proclivity towards a spherical form when suspended in a liquid. At the outset it must have been passive. In respect of its passivity, primitive organic matter must have been like inorganic matter. No such thing as spontaneous variation could have occurred in it; for variation implies some habitual course of change from which it is a divergence, and is therefore excluded where there is no habitual course of change. In the absence of that cyclical series of metamorphoses which even the simplest living thing now shows us, as a result of its inherited constitution, there could be no point d'appui for natural selection. How, then, did organic evolution begin?
If a primitive mass of organic matter was like a mass of inorganic matter in respect of its passivity, and differed only in respect of its greater changeableness; then we must infer that its first changes conformed to the same general law as do the changes of an inorganic mass. The instability of the homogeneous is a universal principle. In all cases the homogeneous tends to pass into the heterogeneous, and the less heterogeneous into the more heterogeneous. In the primordial units of protoplasm, then, the step with which evolution commenced must have been the passage from a state of complete likeness throughout the mass to a state in which there existed some unlikeness. Further, the cause of this step in one of these portions of organic matter, as in any portion of inorganic matter, must have been the different exposure of its parts to incident forces. What incident forces? Those of its medium or environment. Which were the parts thus differently exposed? Necessarily the outside and the inside. Inevitably, then, alike in the organic aggregate and the inorganic aggregate (supposing it to have coherence enough to maintain constant relative positions among its parts), the first fall from homogeneity to heterogeneity must always have been the differentiation of the external surface from the internal contents. No matter whether the modification was physical or chemical, one of composition or of decomposition, it comes within the same generalization. The direct action of the medium was the primordial factor of organic evolution.
In his article on Evolution in the Encyclopædia Britannica, Professor Huxley writes as follows:—
With these passages I may fitly join a remark made in the admirable address Prof. Huxley delivered before unveiling the statue of Mr. Darwin in the Museum at South Kensington. Deprecating the supposition that an authoritative sanction was given by the ceremony to the current ideas concerning organic evolution, he said that "science commits suicide when it adopts a creed."
Along with larger motives, one motive which has joined in prompting the foregoing articles, has been the desire to point out that already among biologists, the beliefs concerning the origin of species have assumed too much the character of a creed; and that while becoming settled they have been narrowed. So far from further broadening that broader view which Mr. Darwin reached as he grew older, his followers appear to have retrograded towards a more restricted view than he ever expressed. Thus there seems occasion for recognizing the warning uttered by Prof. Huxley, as not uncalled for.
Whatever may be thought of the foregoing arguments and conclusions, they will perhaps serve to show that it is as yet far too soon to close the inquiry concerning the causes of organic evolution.
- A Treatise on Comparative Embryology, By F. M. Balfour. Vol. II, chap. xiii.
- Sachs, p. 210.
- Sachs, pp. 83-4.
- Ibid., p. 135.
- Ibid., p. 8.
- Ibid., p. 83.
- Ibid., p. 147.
- A Treatise on Comparative Embryology. By Francis M. Balfour, ll. d., f.r.s. Vol. II, p. 343 (second edition).
- Balfour, l.c. Vol. ii, p. 400-1.
- Ibid., p. 401.
- For a general delineation of the changes by which the development is effected, see Balfour, l.c. Vol. ii, pp. 401-4.