Genesis of Species
Lateral homology (or bilateral symmetry) is the resemblance between the right and left sides of an animal, or of part of an animal; as, e.g., between our right hand and our left. It exists more or less at one or other time of life in all animals, except some very lowly organized creatures. In the highest animals this symmetry is laid down at the very dawn of life, the first trace of the future creature being a longitudinal streak—the embryonic “primitive groove.” This kind of homology is explained by Mr. Spencer as the result of the similar way in which conditions affect the right and left sides respectively.
Vertical homology (or vertical symmetry) is the resemblance existing between parts which are placed one above the other beneath. It is much less general and marked than serial, or lateral homology. Nevertheless, it is plainly to be seen in the tail region of most fishes, and in the far-extending dorsal (back) and ventral (belly) fins of such kinds as the sole and the flounder.
It is also strikingly shown in the bones of the tail of certain efts, as in Chioglossa, where the complexity of the upper (neural) arch is closely repeated by the inferior one. Again, in Spelerpes rubra, where almost vertically ascending articular processes above are repeated by almost vertically descending articular processes below. Also in the axolotl, where there are double pits, placed side by side, not only superiorly but at the same time inferiorly.
This kind of homology is also explained by Mr. Spencer as the result of the similarity of conditions affecting the two parts. Thus he explains the very general absence of symmetry between the dorsal and ventral surfaces of animals by the different conditions to which these two surfaces are respectively exposed, and in the same way he explains the asymmetry of the flat-fishes (Pleuronectidae), of snails, &c.
Now, first, as regards Mr. Spencer’s explanation of animal forms by means of the influence of external conditions, the following observations may be made. Abundant instances are brought forward by him of admirable adaptation of structure to circumstances, but as to the immense majority of these it is very difficult, if not impossible, to see how external conditions can have produced, or even tended to have produced them. For example, we may take the migration of one eye of the sole to the other side of its head. What is there here either in the darkness, or the friction, or in any other conceivable external cause, to have produced the first beginning of such an unprecedented displacement of the eye? Mr. Spencer has beautifully illustrated that correlation which all must admit to exist between the forms of organisms and their surrounding external conditions, but by no means proved that the latter are the cause of the former.
Some internal conditions (or in ordinary language some internal power and force) must be conceded to living organisms, otherwise incident forces must act upon them and upon non-living aggregations of matter in the same way and with similar effects.
If the mere presence of these incident forces produces so ready a response in animals and plants, it must be that there are, in their case, conditions disposing and enabling them so to respond, according to the old maxim, Quicquid recipitur, recipitur ad modum recipientis, as the same rays of light which bleach a piece of silk, blacken nitrate of silver. If, therefore, we attribute the forms of organisms to the action of external conditions, i.e. of incident forces on their modifiable structure, we give but a partial account of the matter, removing a step back, as it were, the action of the internal condition, power, or force which must be conceived as occasioning such ready modifiability. But indeed it is not at all easy to see how the influence of the surface of the ground or any conceivable condition or force can produce the difference which exists between the ventral and dorsal shields of the carapace of a tortoise, or by what differences of merely external causes the ovaries of the two sides of the body can be made equal in a bat and unequal in a bird.
There is, on the other hand, an a priori reason why we should expect to find that the symmetrical forms of all animals are due to internal causes. This reason is the fact that the symmetrical forms of minerals are undoubtedly due to such causes. It is unnecessary here to do more than allude to the beautiful and complex forms presented by inorganic structures. With regard to organisms, however, the wonderful Acanthometrae and the Polycystina may be mentioned as presenting complexities of form which can hardly be thought to be due to other than internal causes. The same may be said of the great group of Echinoderms, with their amazing variety of component parts. If then internal forces can so build up the most varied structures, they are surely capable of producing the serial, lateral, and vertical symmetries which higher animal forms exhibit. Mr. Spencer is the more bound to admit this, inasmuch as in his doctrine of “physiological units” he maintains that these organic atoms of his have an innate power of building up and evolving the whole and perfect animal from which they were in each case derived. To build up and evolve the various symmetries here spoken of is not one whit more mysterious. Directly to refute Mr. Spencer’s assertion, however, would require the bringing forward of examples of organisms which are ill-adapted to their positions, and out of harmony with their surroundings—a difficult task indeed.
Secondly, as regards the last-mentioned author’s explanation of such serial homology as exists in the centipede and its allies, the very groundwork is open to objection. Multiplication by spontaneous fission seems from some recent researches to be much less frequent than has been supposed, and more evidence is required as to the fact of the habitual propagation of any planariae in this fashion. But even if this were as asserted, nevertheless it fails to explain the peculiar condition presented by Syllis and some other annelids, where a new head is formed at intervals in certain segments of the body. Here there is evidently an innate tendency to the development at intervals of a complex whole. It is not the budding out or spontaneous fission of certain segments, but the transformation in a definite and very peculiar manner of parts which already exist into other and more complex parts. Again, the processes of development presented by some of these creatures do not by any means point to an origin through the linear coalescence of primitively distinct animals by means of imperfect segmentation. Thus in certain Diptera (two winged flies) the legs, wings, eyes, &c., are derived from masses of formative tissue (termed imaginal disks), which by their mutual approximation together build up parts of the head and body, recalling to mind the development of Echinoderms.
Again, Nicholas Wagner found in certain other Diptera, the Hessian flies, that the larva gives rise to secondary larvae within it, which develop and burst the body of the primary larva. The secondary larvae give rise, similarly, to another set within them, and these again to another set.
Again, the fact that in Taenia echinococcus one egg produces numerous individuals, tends to invalidate the argument that the increase of segments during development is a relic of specific genesis.
Mr. H. Spencer seems to deny serial homology to the mollusca, but it is difficult to see why the shell segments of chiton are not such homologues because the segmentation is superficial. Similarly the external processes of eolis, doris, &c., are good examples of serial homology, as also are plainly the successive chambers of the orthoceratidae. Nor are parts of a series less serial, because arranged spirally, as in most gasteropods. Mr. Spencer observes of the molluscous as of the vertebrate animal, “You cannot cut it into transverse slices, each of which contains a digestive organ, a respiratory organ, a reproductive organ, &c.” But the same may be said of every single arthropod and annelid if it be meant that all these organs are not contained in every possible slice. While if it be meant that parts of all such organs are contained in certain slices, then some of the mollusca may also be included.
Another objection to Mr. Spencer’s speculation is derived from considerations which have already been stated, as to past time. For if the annulose animals have been formed by aggregation, we ought to find this process much less perfect in the oldest form. But a complete development, such as already obtains in the lobster, &c., was reached by the Eurypterida and Trilobites of the palaeozoic strata; and annelids, probably formed mainly like those of the present day, abounded during the deposition of the oldest fossiliferous rocks.
Thirdly, and lastly, as regards such serial homology as is exemplified by the backbone of man, there are also several objections to Mr. Spencer’s mechanical explanation.
On the theory of evolution most in favour, the first Vertebrata were aquatic. Now, as natation is generally effected by repeated and vigorous lateral flexions of the body, we ought to find the segmentation much more complete laterally than on the dorsal and ventral aspects of the spinal column. Nevertheless, in those species which, taken together, constitute a series of more and more distinctly segmented forms, the segmentation gradually increases all round the central part of the spinal column.
Mr. Spencer thinks it probable that the sturgeon has retained the notochordal (that is, the primitive, unsegmented) structure because it is sluggish. But Dr. Guenther informs me that the sluggishness of the common tope (Galeus vulgaris) is much like that of the sturgeon, and yet the bodies of its vertebrae are distinct and well-ossified. Moreover, the great salamander of Japan is much more inert and sluggish than either, and yet it has a well-developed, bony spine.
I can learn nothing of the habits of the sharks Hexanchus, Heptanchus, and Echinorhinus, but Mueller describes them as possessing a persistent _chorda dorsalis_. It may be they have the habits of the tope, but other sharks are amongst the very swiftest and most active of fishes.
In the bony pike (lepidosteus), the rigidity of the bony scales by which it is completely enclosed must prevent any excessive flexion of the body, and yet its vertebral column presents a degree of ossification and vertebral completeness greater than that found in any other fish whatever.