Francis M. Balfour

“ If a naturalist were to study the life-history, or, in other words, the embryology of this form, this fact about the pigment-spot would come to his notice, and he would be justified, from the laws of heredity, in concluding that the species was descended from an ancestor without a pigment-spot, because a pigment-spot was absent in the young. Now, we may suppose the transparent layer of skin above the pigment-spot to become thickened, so as gradually to form a kind of lens, which would throw an image of external objects on the pigment-spot. ”

“ They exhibit a division of labour: some carrying on the work of digestion; some, which we call nerve-cells, receiving and conducting stimuli; some, which we call muscle-cells, altering their form—in fact, contracting in one direction—under the action of the stimuli brought to them by the nerve-cells. In most cases a number of cells with the same function are united together, and thus constitute a tissue. Thus the cells which carry on the work of digestion form a lining membrane to a tube or sack, and constitute a tissue known as a secretory epithelium. ”

“ As development proceeds the vesicle of the hemispheres becomes divided by the ingrowth of a median longitudinal septum, and the olfactory lobes grow out from the posterior lateral regions of each ventricle thus formed, and eventually rise on to the dorsal faces of the hemispheres, instead of, as in most Vertebrata, remaining on their ventral sides. I may remark, that I cannot accept the views of Miklucho-Maclay, whose proposal to alter the nomenclature of the parts of the Elasmobranch's brain, appears to me to be based upon a misinterpretation of the facts of development. ”

“ It is a point of some importance for understanding the relation between the mode of formation of the alimentary canal in the Frog and other vertebrates to notice that on the ventral surface the cells which are to form the epithelium of the alimentary canal become distinguished as such very much later than do those to form its dorsal epithelium, and are derived not from the involuted cells but from the primitive large yolk-cells. It is indeed probable that only a very small portion of epithelium of the ventral wall of the mid-gut is in the end derived from these larger yolk-cells. ”

“ The most important part of the central nervous system of these animals is the supra-œsophageal ganglia, often known as the brain, and these are formed in the embryo from two thickened patches of the skin at the front end of the body. These thickened patches become gradually detached from the surface, remaining covered over by a layer of skin. They then constitute the supra-œsophageal ganglia; but they form not only the ganglia, but also the rhabdons or retinal elements of the eye—the parts in fact which correspond to the rods and cones in our own retina. ”

“ As there, the primitive involution is merely represented by a split separating the yolk and the germinal disc, and on the dorsal side alone is there a true cellular wall for this split, and at the dorsal mouth of the split the alimentary epithelium becomes continuous with the epiblast.
The segmentation cavity has become diminished, and round the yolk the epiblast, accompanied by a layer of mesoblast, is commencing to grow. In this growth all parts of the blastoderm take a share except that part where the epiblast and hypoblast are continuous. ”

“ In this case we would be compelled to suppose that the mass of lower layer cells which forms the embryonic swelling is used as food for the growth of the involuted epiblast, or else employed solely in the growth over the yolk of the non-embryonic portion of the blastoderm; but the latter possibility does not seem compatible with my sections.
I do not believe that it is possible, from the examination of sections alone, to decide which of these two views (viz. whether the epiblast is involuted, or whether it becomes merely continuous with the lower layer cells) is the true one. ”

“ Secondly, the body-cavity when it appears in the mesoblast plates, does not arise as a single cavity, but as a pair of cavities, one for each plate of mesoblast, and these cavities remain permanently distinct in some parts of the body, and nowhere unite till a comparatively late period. Thirdly, the primitive body-cavity of the embryo is not confined to the region in which a body-cavity exists in the adult, but extends to the summit of the muscle-plates, at first separating parts which become completely fused in the adult to form the great lateral muscles of the body. ”

“ Had Balfour been only an ordinary man, the zeal and energy which he threw into his lectures, and into the supervision of the practical work, added to the almost brotherly interest which he took in the individual development of every one of the pupils who shewed any love whatever for the subject, would have made him a most successful teacher. ”

“ As growth continues, the primitive groove becomes pushed further and further back, and becomes less and less conspicuous, till at about thirty-six hours only a very small and curved remnant is to be seen behind the sinus rhomboidalis; but even up to the forty-ninth Dursy has been able to distinguish it at the hinder end of the embryo.
The primitive groove in the chick is, then, a structure which appears very early, and soon disappears without entering directly into the formation of any part of the future animal, and without, so far as I can see, any function whatever. ”

“ The tissue of the neural arches does not any longer form a continuous sheet, but is divided into (1) a series of arches encircling the spinal cord, and (2) a basal portion resting on the cartilaginous sheath of the notochord. There are two arches to each muscle-plate, one continuous with the basal portion of the arch-tissue and forming the true arch, which springs opposite the centre of a vertebral body, and the second not so continuous, which forms what is usually known as the intercalated piece. Between every pair of true arches the two roots of a single spinal nerve pass out. ”

“ The body is always built of certain pieces of protoplasm, which are technically known to biologists as cells. The simplest organisms are composed either of a single piece of this kind, or of several similar pieces loosely aggregated together. Each of these pieces or cells is capable of digesting and assimilating food, and of respiring; it can execute movements, and is sensitive to external stimuli, and can reproduce itself. All the functions of higher animals can, in fact, be carried on in this single cell. Such lowly organized forms are known to naturalists as the Protozoa. ”

“ It is, indeed, very easy to see that most freshly formed ova are derived from nests, as in the embryo; and the formation of a follicular epithelium round these ova takes place as they become separated from the nests. A few solitary ova, which have never formed part of a nest, seem to be formed in this stage as in the embryo; but they do not grow into the stroma surrounded by the cells of the pseudo-epithelium, and only as they reach a not inconsiderable size is a definite follicular epithelium formed around them. ”

“ As has already been pointed out the transformation which the so-called primitive ova undergo is sufficient to shew that they are not to be regarded as ova but merely as embryonic sexual cells. A feature in the transformation, which appears to be fairly constant in Scyllium, and not uncommon in the rabbit, is the fusion of the protoplasm of several ova into a syncytium, the subsequent increase in the number of nuclei in the syncytium, the atrophy and absorption of a portion of the nuclei, and the development of the remainder into the germinal vesicles of ova ”

“ The cranial flexure, which commenced in stage G, has now become very evident, and the mid-brain [178] begins to project in the same manner as in the embryo fowl on the third day, and will soon form the anterior termination of the long axis of the embryo. The fore-brain has increased in size and distinctness, and the anterior part of it may now be looked on as the unpaired rudiment of the cerebral hemispheres.
Further growths have taken place in the organs of sense, especially in the eye, in which the involution for the lens has made considerable progress. ”

“ If such points as these can be affected by natural selection I see no reason why the arrangement of individual cells (or rather primitive elements) should not also be modified; why, in fact, a mass of cells which was originally derived from one layer, but in the course of development became budded off from that layer and entered another layer, should not by a series of small steps cease ever to be attached to the original layer, but from the first moment it can be distinguished should be found as a separate mass in the second layer. ”

“ In so excessively highly differentiated a type as birds we could hardly expect to find, and certainly do not find, any traces of the [Pg 35] primitive origin of the mesoblast, either from the epiblast or hypoblast, or from both. In the chick the mesoblast cells are formed directly from the ultimate products of segmentation. From having a secondary origin in most invertebrates the mesoblast comes to have, in the chick, a primary origin from the segmentation spheres, precisely as we find to be the case with the nervous layer in osseous fishes. ”

“ It will hardly be disputed that primitively the muscular system of the body-wall could not have been derived from the layer of cells which lines the alimentary canal. We see indeed in Hydra and the Hydrozoa that in its primitive differentiation, as could have been anticipated beforehand, the muscular system of the body is derived from the epiblast cells. What, then, is the explanation of the widespread derivation of the mesoblast, including the muscular system of the body, from the hypoblast? ”

“ In the first place every Metazoon commences its existence as a simple cell, in the sense above defined; this cell is known as the ovum. The first developmental process which takes [Pg 704] place consists in the division or segmentation of the single cell into a number of smaller cells. The cells then arrange themselves into two groups or layers known to embryologists as the primary germinal layers. These two layers are usually placed one within the other round a central cavity. The inner of the two is called the hypoblast, the outer the epiblast. ”

“ The heart has now, in most respects, reached its full development. It is formed of an outer muscular layer, within which is a doubly-contoured lining, containing nuclei at intervals, which is probably of the nature of an epithelioid lining (Pl. 32, fig. 22, ht) . In its lumen are numerous blood-corpuscles (not represented in my figure) . The heart lies in a space bound below by the splanchnic mesoblast, and to the sides by the somatic mesoblast. This space forms a kind of pericardium (fig. 22, pc) , but dorsally the heart is in contact with the epiblast. ”