embryos of closely allied animals are distinguishable at all stages of development, though the distinguishing features are not the same as those which distinguish the adults. To say that the development of the organism and of its component parts is a progress from the simple to the complex is to state a truism, but to state that it is also a progress from the general to the special is to go altogether beyond the facts. The bipinnaria larva of an echinoderm, the trochosphere larva of an annelid, the blastodermic vesicle of a mammal are all as highly specialized as their respective adults, but the specialization is for a different purpose, and of a different kind to that which characterizes the adult.
In its scientific and systematic form embryology may be considered as having only taken birth within the last century, although the germ from which it sprung was already formed nearly half a century earlier. The ancients, it is true, as we see by the writings of Aristotle and History of embryology. Galen, pursued the subject with interest, and the indefatigable Greek naturalist and philosopher had even made continued series of observations on the progressive stages of development in the incubated egg, and on the reproduction of various animals; but although, after the revival of learning, various anatomists and physiologists from time to time made contributions to the knowledge of the foetal structure in its larger organs, yet from the minuteness of the observations required for embryological research, it was not till the microscope came into use for the investigation of organic structure that any intimate knowledge was attained of the nature of organogenesis. It is not to be wondered at, therefore, that during a long period, in this as in other branches of physical inquiry, vague speculations took the place of direct observation and more solid information. This is apparent in most of the works treating of generation during the 16th and part of the 17th centuries.[1]
Harvey was the first to give, in the middle of the latter century, a new life and direction to investigation of this subject, by his discovery of the connexion between the cicatricula of the yolk and the rudiments of the chick, and by his faithful description of the successive stages of development as observed in the incubated egg, as well as of the progress of gestation in some Mammalia. He had also the merit of fixing the attention of physiologists upon general laws of development as deduced from actual observation of the phenomena, by the enunciation of two important propositions, viz.—(1) that all animals are produced out of ova, and (2) that the organs of the embryo arise by new formation, or epigenesis, and not by mere enlargement out of a pre-existing invisible condition (Exercitationes de generatione animalium, Amstelodami, 1651). Harvey’s observations, however, were aided only by the use of magnifying glasses (perspecillae), probably of no great power, and he saw nothing of the earliest appearances of the embryo in the first thirty-six hours, and believed the blood and the heart to be the parts first formed.
The influence of the work of Harvey, and of the successful application of the microscope to embryological investigation, was soon afterwards apparent in the admirable researches of Malpighi of Bologna, as evinced by his communications to the Royal Society of London in 1672, “De ovo incubato,” and “De formatione pulli,” and more especially in his delineations of some of the earlier phenomena of development, in which, as in many other parts of minute anatomy, he partially or wholly anticipated discoveries, the full development of which has only been accomplished in the present century. Malpighi traced the origin of the embryo almost to its very commencement in the formation of the cerebro-spinal groove within the cicatricula, which he removed from the opaque mass of the yolk; and he only erred in supposing the embryonal rudiments to have pre-existed as such in the egg, in consequence, apparently, of his having employed for observation, in very warm weather, eggs which, though he believed them to be unincubated, had in reality undergone some of the earlier developmental changes.
The works of Walter Needham (1667), Regnier de Graaf (1673), Swammerdam (1685), Vallisneri (1689)—following upon those of Harvey—all contain important contributions to the knowledge of our subject, as tending to show the similarity in the mode of production from ova in a variety of animals with that previously best known in birds. The observations more especially of de Graaf, Nicolas Steno and J. van Horne gave much greater precision to the knowledge of the connexion between the origin of the ovum of quadrupeds and the vesicles of the ovary now termed Graafian, which de Graaf showed always burst and discharged their contents on the occurrence of pregnancy.
These observations bring us to the period of Boerhaave and Albinus in the earlier part of the 18th century, and in the succeeding years to that of Haller, whose vast erudition and varied and accurate original observations threw light upon the entire process of reproduction in animals, and brought its history into a more systematic and intelligible form. A considerable part of the seventh and the whole of the eighth volumes of Haller’s great work, the Elementa physiologiae, published at successive times from 1757 to 1766, are occupied with the general view of the function of generation, while his special contributions to embryology are contained in his Deux mémoires sur la formation du cœur dans le poulet and Deux mémoires sur la formation des os, both published at Lausanne in 1758, and republished in an extended and altered form, together with his “Observations on the early condition of the Embryo in Quadrupeds,” made along with Kühlemann, in the Opera minora (1762–1768). Though originally educated as a believer in the doctrine of “preformation” by his teacher Boerhaave, Haller was soon led to abandon that view in favour of “epigenesis” or new formation, as may be seen in various parts of his works published before the middle of the century; see especially a long note explanatory of the grounds of his change of opinion in his edition of Boerhaave’s Praelectiones academicae, vol. v. part 2, p. 497 (1744), and his Primae lineae physiologiae (1747). But some years later, and after having been engaged in observing the phenomena of development in the incubated egg, he again changed his views, and during the remainder of his life was a keen opponent of the system of epigenesis, and a defender and exponent of the theory of “evolution,” as it was then named—a theory very different from that now bearing the name, and which implied belief in the pre-existence of the organs of the embryo in the germ, according to the theory of encasement (emboîtement) or inclusion supported by Leibnitz and Bonnet. (See the interesting work of Bonnet, Considérations sur les corps organisés, Amsterdam, 1762, for an account of his own views and those of Haller.)
It was reserved for Caspar Frederick Wolff (1733–1794), a German by birth, but naturalized afterwards in Russia, to bring forward observations which, though almost entirely neglected for a long time after their publication, and in some measure discredited under the influence of Haller’s authority, were sixty years later acknowledged to have established the theory of epigenesis upon the secure basis of ascertained facts, and to have laid the first foundation of the morphological science of embryology. Wolff’s work, entitled Theoria generationis, first published as an inaugural Dissertation at Berlin in 1759, was republished with additions in German at Berlin in 1764, and again in Latin at Halle in 1774. Wolff also wrote a “Memoir on
- ↑ It may be proper to mention, as authors of this period who made special researches on the development of the embryo—(1) Volcher Coiter of Groningen, who, along with Aldrovandus of Bologna, made a series of observations on the formation of the chick, day by day, in the incubated egg, which were described in a work published in 1573, and (2) Hieronymus Fabricius (ab Aquapendente), who, in his work De formato foetu, first published at Padua in 1600, gave an interesting account, illustrated by many fine engravings, of uterogestation and the foetus of a number of quadrupeds and other animals, and in a posthumous work entitled De formatione ovi et pulli, edited by J. Prevost and published at Padua in 1621, described and illustrated by engravings the daily changes of the egg in incubation. It is enough, however, to say that Fabricius was entirely ignorant of the earlier phenomena of development which occur in the first two or three days, and even of the source of the embryonic rudiments, which he conceived to spring, not from the yolk or true ovum, but from the chalazae or twisted, deepest part of the white. The cicatricula he looked upon as merely the vestige of the pedicle by which the yolk had previously been attached to the ovary.
