[4] The value of these identifications as well as of those below is discussed in its appropriate place in the body of the work. Their citation here is not to be regarded as necessarily implying my acceptance of them.
(2) How far some special embryonic larval form is constantly reproduced in the ontogeny of the members of one or more groups of the animal kingdom; and how far such larval forms may be interpreted as the ancestral type for those groups.
As examples of such forms may be cited the six-limbed Nauplius supposed by Fritz Müller to be the ancestral form of the Crustacea; the trochosphere larva of Lankester, which he considers to be common to the Mollusca, Vermes, and Echinodermata; the planula of the Coelenterata, etc.
(3) How far such forms agree with living or fossil forms in the adult state; such an agreement being held to imply that the living or fossil form in question is closely related to the parent stock of the group in which the larval form occurs. It is not easy to cite examples of a very close agreement of this kind between the larval forms of one group and the existing or fossil forms of another. The larvæ of some of the Chætopoda with long provisional setæ resemble fossil Chætopods. The Rotifers have many points of resemblance to the trochosphere, especially to that form of trochosphere characteristic of the Mollusca. The Turbellarians have some features in common with the Coelenterate planula. Some of the Gephyrea in the presence of a præoral lobe resemble certain trochosphere types. The larva of the Tunicata has the characters of a simple type of the Chordata.
Within the limits of a single group agreements of this kind are fairly numerous. In the Craniata the tadpole of the Anura has its living representative in the Pisces and perhaps especially in the Myxinoids. The larval forms of the Insecta approach Peripatus. The stalked larva of Comatula is reproduced by the living Pentacrinus and Rhizocrinus etc. Numerous examples of the same phenomenon are found amongst the Crustacea.
(4) How far organs appear in the embryo or larva which either atrophy or become functionless in the adult state, and which persist permanently in members of some other group or in lower members of the same group. Cases of this kind are of the most constant occurrence, and it is only necessary to cite such examples as the gill slits and Wolffian body in the embryos of higher Craniata to illustrate the kind of instance alluded to. The same conclusions may be drawn from them as from the cases under the previous heading.
(5) How far organs pass in the course of their development through a condition permanent in some lower form. Phylogenetic conclusions may be drawn from instances of this character, though they have a more important bearing on Organology than on Phylogeny.
The considerations which were used to shew that the ancestral history is reproduced in the ontogeny of the individual apply with equal force to the evolution of organs. The special questions in Organology, on which Comparative Embryology throws light, may be classified under the following heads.
(1) The origin and homologies of what are known as the germinal layers; or the layers into which the embryo becomes divided immediately after the segmentation.
(2) The origin of primary tissues, epithelial, nervous, muscular, connective, etc., and their relation to the germinal layers.
(3) The origin of organs. The origin of the primitive organs is intimately connected with that of the germinal layers. The first differentiation of the segmented ovum results in the cells of the embryo becoming arranged as two layers, an outer one known as the epiblast and an inner one as the hypoblast. The outer of these forms a primitive sensory organ, and the inner a primitive digestive organ.
(4) The gradual evolution of the more complicated organs and systems of organs.
This part of the subject, even more than that dealing with questions of Phylogeny, is intimately bound up with Comparative Anatomy; without which indeed it becomes quite meaningless.
REPRODUCTION.
A study of reproduction logically precedes that of Embryology. Reproduction essentially consists in the separation of a portion of an organism which has the capacity of developing into a form similar to that which gave it origin. The simplest modes of reproduction are those which occur amongst the Protozoa.
In this group, reproduction may take place in a great variety of ways. These may be classified in three groups: (1) fission, (2) budding or gemmation, (3) spore formation.
Reproduction in all these ways may take place either subsequently to and apparently in consequence of a very important process known as conjugation, which consists in the temporary or permanent fusion of two or more individuals, or spontaneously, i.e. independently of any such previous conjugation.
Reproduction by fission consists simply in the division of the organism into two similar parts, the nucleus when present becoming divided simultaneously with the cell body. This mode of reproduction is the simplest conceivable, and is not followed by a development, since the two organisms produced are exactly similar, except in size, to the parent form. Besides single fission, a process of multiple fission may take place, as amongst the Flagellata, where Drysdale and Dallinger have shewn that an individual enclosed within a structureless cyst may divide first into two, then into four, and so on.
The process of budding differs mainly from that of simple fission in the fact that the two organisms produced are dissimilar in size, and also that the separation of the smaller organism from the larger is preceded by a process of growth in the latter, so that in the separation of the bud no essential part of the parent form is removed. This mode of reproduction is found amongst the Infusoria, Acineta, &c. An interesting variation in it is the internal gemmation of many of the Acineta, where a portion of the internal protoplasm with part of the nucleus is separated off to form a fresh individual. This mode of gemmation is connected by a series of gradations with the normal external gemmation. The organisms produced by gemmation are not always similar at birth to the parent; e.g. Acineta.
Both fission and gemmation when incomplete lead to the formation of colonies.
The third mode of reproduction, by spore formation, does not essentially differ from that by multiple fission. It consists in the breaking up of the organisms into a number (usually very considerable) of portions; each of which eventually develops into an organism like the parent form. All gradations between a simultaneous division of the organism into such spores and simple multiple fission are to be found, but this process of reproduction may be sometimes distinguished from that by such fission by the fact that the two processes may coexist in a single form, e.g. the biflagellate monad of Drysdale and Dallinger. In the majority of cases the spores produced differ at first from the parent organism not only in size but in other points, such as the possession of a flagellum, etc. They may even be without a nucleus when the parent organism is nucleated, as in the Gregarinidæ.
The encystment, which in many cases precedes reproduction by any of the above processes, and more especially by spores, is not an essential condition of their occurrence; and is probably in the first instance a protective arrangement which has become secondarily adapted to and connected with reproduction.
As has been already stated, all the above modes of reproduction take place in some of the Protozoa without any anterior process which can be regarded as of a sexual nature; but very often they are preceded by the temporary or permanent fusion of two or more individuals, such fusion being known as conjugation.
In most cases reproduction by spores is the consequence of conjugation, but in the Infusoria etc. where the fusion at conjugation is temporary (except Vorticella), there is probably merely a renewed activity--a rejuvenescence--which most likely results in active fission or budding. In the Gregarinidæ reproduction by spores usually follows conjugation, but may also take place without it. In some Flagellata reproduction by spores follows the conjugation of two individuals in a different stage of development. Thus in the springing Monad, described by Drysdale and Dallinger, a form produced by the fission of a monad in an amoeboid condition fuses with an ordinary monad to produce an individual, which then breaks up into spores. Another instance of the fusion of dissimilar individuals is afforded by Vorticella, where a free-swimming individual conjugates and is permanently united with a fixed one (Engelmann, Bütschli). Conjugation often consists in the fusion of more than two individuals. In conjugation where the fusion is permanent, the nuclei of the conjugating forms usually unite before the product breaks up into spores and where temporary fusion occurs in the Infusoria a division of the paranuclei and often of the nuclei takes place, followed by the ejection of parts of them, and a reproduction of new paranuclei and nuclei from the remainder of the original structures.