Echinoderms (Hensen, Agassiz, Metschnikoff, Selenka, Götte), Nematodes (Bütschli), Sagitta (Kowalevsky, Bütschli), Lumbricus and probably other Annelids (Kowalevsky), Brachiopoda (Kowalevsky), Crustaceans (Bobretzky), Insects (Kowalevsky, Ulianin, Dohrn), Myriapods (Metschnikoff), Tunicates (Kowalevsky, Kuppfer), Petromyzon (Owsjanikoff), Osseous fishes (Oellacher, Götte, Kowalevsky), Elasmobranchii (Self), Amphibians (Remak, Stricker, Götte).
The list includes members from the greater number of the groups of the animal kingdom; the most striking omissions being the Cœlenterates, Mollusks, and the Amniotic Vertebrates. The absence of the Cœlenterates has been already explained and my grounds for regarding the Amniotic Vertebrates as apparent rather than real exceptions have also been pointed out. The Mollusks, however, remain as a large group, in which we as yet know very little as to the formation of the mesoblast.
Dr Rabl[216], who seems recently to have studied the development of Lymnæus by means of sections, gives some figures shewing the origin of the mesoblast; they are, however, too diagrammatic to be of much service in settling the present question, and the memoirs of Professor Lankester[217] and Dr Fol[218] are equally inconclusive for this purpose, for, though they contain figures of elongated and branched mesoblast cells passing from the epiblast to the hypoblast, no satisfactory representations are given of the origin of these cells. I have myself observed in embryos of Turbo or Trochus similar elongated cells to those figured by Lankester and Fol, but was unable clearly to determine whence they arose. The most accurate observations which we have on this question are those of Professor Bobretzky[219]. In Nassa he finds that the three embryonic layers are all established during segmentation. The outermost and smallest cells form the epiblast, somewhat larger cells adjoining these the mesoblast, and the large yolk-cells the hypoblast. These observations do not, however, demonstrate from which of the primary layers the mesoblast is derived.
The evidence at present existing is clearly in favour of the mesoblast being, in almost all groups of animals, developed from the hypoblast, but strong as this evidence is, it has not its full weight unless the actual manner in which the mesoblast is in many groups derived from the hypoblast, is taken into consideration. The most important of these are the Echinoderms, Brachiopods and Sagitta.
In the Echinoderms the mesoblast is in part formed by cells budded off from the hypoblast, the remainder, however, arises as one or more diverticula of the alimentary tract. From the separate cells first budded off there are formed the cutis, part of the connective tissue and the calcareous skeleton[220]. The diverticula from the alimentary cavity form the water-vascular system and the somatic and splanchnic layers of mesoblast. The cavity of the diverticula after the separation of the water-vascular system, forms the body-cavity. The outer lining layer of the cavity forms the somatic layer of mesoblast and the voluntary muscles; the inner lining layer the splanchnic mesoblast which unites with the epithelium of the alimentary tract. Though this fundamental arrangement would seem to be universal amongst Echinoderms, considerable variations of it are exhibited in different groups.
There is one outgrowth from the alimentary tract in Synapta; two in Echinoids, Asteroids and Ophiura; three in Comatula, and four (?) in Amphiura. The cavity of the outgrowth usually forms the body-cavity, but sometimes in Ophiura and Amphiura (Metschnikoff) the outgrowths are from the first or soon become solid, and only secondarily acquire a cavity, which is however homologous with the body-cavity of the other groups.
In Sagitta[221] the formation of the mesoblast and the alimentary tract takes place in nearly the same fashion as in the Echinoderms. The simple invaginate alimentary cavity becomes divided into three lobes, a central and two lateral. The two lateral lobes are gradually more and more constricted off from the central one, and become eventually quite separated from it; their cavities remain independent, and form in the adult the body-cavity, divided by a mesentery into two distinct lateral sections. The inner layer of each of the two lateral lobes forms the mesoblast of the splanchnopleure, the outer layer the mesoblast of the somatopleure. The central division of the primitive gastræa cavity remains as the alimentary tract of the adult.
The remarkable observations of Kowalevsky[222] on the development of the Brachiopoda have brought to light the unexpected fact that in two genera at least (Argiope and Terebratula) the mesoblast and body-cavity develop as paired constrictions from the alimentary tract in a manner almost identically the same as in Sagitta.
It thus appears that, so far as can be determined from the facts at our disposal, the mesoblast in almost all cases is derived from the hypoblast, and in three widely separated groups it arises as a pair of diverticula from the alimentary tract, each diverticulum containing a cavity which eventually becomes the body-cavity. I have elsewhere suggested[223] that the origin of the mesoblast from alimentary diverticula is to be regarded as primitive for all higher animals, and that the more general cases in which the mesoblast becomes split off, as an undivided layer, from the hypoblast, are in reality derivates from this. The chief obstacle in the way of this view arises from the difficulty of understanding how the whole voluntary muscular system can have been derived at first from the alimentary tract. That part of a voluntary system of muscles might be derived from the contractile diverticula of the alimentary canal attached to the body-wall is not difficult to understand, but it is not easy to believe that the secondary system so formed could completely replace the primitive muscular system, derived, as it must have been, from the epiblast. In my paper above quoted will be found various speculative suggestions for removing this difficulty, which I do not repeat here. If it be granted, however, that in Sagitta, Brachiopods, and Echinoderms we have genuine examples of the formation of the whole mesoblast from alimentary diverticula, it is easy to see how the formation of the mesoblast in Vertebrates may be a secondary derivate from an origin of this nature.
An attempt has been already made to shew that the mesoblast in Elasmobranchii is formed in a very primitive fashion, and for this reason the Elasmobranchii appear to be especially adapted for determining whether any signs are exhibited of a derivation of the mesoblast as paired diverticula of the alimentary tract. There are, it appears to me, several such features. In the first place, the mesoblast is split off from the hypoblast not as a single mass but as a pair of distinct masses, comparable with the paired diverticula already alluded to. 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. It is difficult to understand how the body-cavity could have such an extension as this, on the supposition that it represents a primitive split in the mesoblast between the wall of the gut and the body-wall; but its extension to this part is quite intelligible, on the supposition that it represents the cavities of two diverticula of the alimentary tract, from whose muscular walls the voluntary muscular system has been derived. Lastly, I would point out that the derivation of part of the muscular system from what appears as the splanchnopleure is quite intelligible on the assumed hypothesis, but, as far as I see, on no other.