THE WORKS

OF

FRANCIS MAITLAND BALFOUR.

VOL. II.

Memorial Edition.

Cambridge:

PRINTED BY C. J. CLAY, M.A. AND SON,
AT THE UNIVERSITY PRESS.

Memorial Edition.

THE WORKS
OF
FRANCIS MAITLAND BALFOUR,

M.A., LL.D., F.R.S.,

FELLOW OF TRINITY COLLEGE,
AND PROFESSOR OF ANIMAL MORPHOLOGY IN THE UNIVERSITY OF
CAMBRIDGE.

EDITED BY

M. FOSTER, F.R.S.,
PROFESSOR OF PHYSIOLOGY IN THE UNIVERSITY OF CAMBRIDGE;

AND

ADAM SEDGWICK, M.A.,
FELLOW AND LECTURER OF TRINITY COLLEGE, CAMBRIDGE.

VOL. II.

A TREATISE ON COMPARATIVE EMBRYOLOGY.

Vol. I. Invertebrata.

London:
MACMILLAN AND CO.
1885

[The Right of Translation is reserved.]

PREFACE.

My aim in writing this work has been to give such an account of the development of animal forms as may prove useful both to students and to those engaged in embryological research. The present volume, save in the introductory chapters, is limited to a description of the development of the Invertebrata: the second and concluding volume will deal with the Vertebrata, and with the special histories of the several organs.

Since the work is, I believe, with the exception of a small but useful volume by Packard, the first attempt to deal in a complete manner with the whole science of Embryology in its recent aspects, and since a large portion of the matter contained in it is not to be found in the ordinary text books, it appeared desirable to give unusually ample references to original sources. I have accordingly placed at the end of each chapter, or in some cases of each section of a chapter, a list of the more important papers referring to the subject dealt with. The papers in each list are numbered continuously, and are referred to in the text by their numbers. These lists are reprinted as an appendix at the end of each volume. It will of course be understood that they do not profess to form a complete bibliography of the subject.

In order to facilitate the use of the work by students I have employed two types. The more general parts of the work are printed in large type; while a smaller type is used for much of the theoretical matter, for the details of various special modes of development, for the histories of the less important forms, and for controversial matter generally. The student, especially when commencing his studies in Embryology, may advantageously confine his attention to the matter in the larger type; it is of course assumed that he already possesses a competent knowledge of Comparative Anatomy.

Since the theory of evolution became accepted as an established doctrine, the important bearings of Embryology on all morphological views have been universally recognised: but the very vigour with which this department of science has been pursued during the last few years has led to the appearance of a large number of incomplete and contradictory observations and theories; and to arrange these into anything like an orderly and systematic exposition has been no easy task. Many Embryologists will indeed probably hold that any attempt to do so at the present time is premature, and therefore doomed to failure. I must leave it to others to decide how far my effort has been justified. That what I have written contains errors and shortcomings is I fear only too certain, but I trust that those who are most capable of detecting them will also be most charitable in excusing them.

The work is fully illustrated, and most of the figures have been especially engraved from original memoirs or from my own papers or drawings by Mr Collings, who has spared no pains to render the woodcuts as clear and intelligible as possible. I trust my readers will not be disappointed with the results. The sources from which the woodcuts are taken have been in all cases acknowledged, and in the cases where no source is given the illustrations are my own.

I take this opportunity of acknowledging my great obligations to Professors Agassiz, Huxley, Gegenbaur, Lankester, Turner, Kölliker, and Claus, to Sir John Lubbock, Mr Moseley, and Mr P. H. Carpenter, for the use of electrotypes of woodcuts from their works.

I am also under great obligations to numerous friends who have helped me in various ways in the course of my labour. Professor Kleinenberg, of Messina, has read through the whole of the proofs, and has made numerous valuable criticisms. My friend and former pupil, Mr Adam Sedgwick, has been of the greatest assistance to me in correcting the proofs. I have had the benefit of many useful suggestions by Professor Lankester especially in the chapter on the Mollusca, and Mr P. H. Carpenter has kindly revised the chapter on the Echinodermata.

I am also much indebted to Dr Michael Foster, Mr Moseley, and Mr Dew-Smith for aid and advice.

CONTENTS OF VOLUME I.

Introduction. Pp. [1‑16].

Chapter I. The Ovum and Spermatozoon.

General history of the Ovum, pp. [17‑25]. Special history of the Ovum in different types, pp. [26‑65]. The Spermatozoon, pp. [65‑67].

Chapter II. The Maturation and Impregnation of the Ovum.

Maturation of the Ovum, and formation of the polar bodies, pp. [68‑79]. Impregnation of the Ovum, pp. [79‑86]. Summary, p. [86].

Chapter III. The Segmentation of the Ovum.

Internal phenomena of Segmentation, pp. [88‑92]. External features of Segmentation, pp. [92‑122].

Introduction to Systematic Embryology. Pp. [125‑130].

Chapter IV. Dicyemidæ and Orthonectidæ. Pp. [131‑137].

Chapter V. Porifera. Pp. [138‑151].

Chapter VI. Cœlenterata.

Hydrozoa, pp. [152‑167]. Actinozoa, pp. [167‑173]. Ctenophora, pp. [173‑178]. Summary, etc., pp. [178‑182]. Alternations of generations, pp. [182‑187].

Chapter VII. Platyelminthes.

Turbellaria, pp. [189‑196]. Nemertea, pp. [196‑204]. Trematoda, pp. [205‑210]. Cestoda, pp. [210‑218].

Chapter VIII. Rotifera. Pp. [221‑224].

Chapter IX. Mollusca.

Formation of the layers and larval characters, pp. [225‑273]. Gasteropoda and Pteropoda, pp. [225‑242]. Cephalopoda, pp. [242‑254]. Polyplacophora, pp. [254‑257]. Scaphopoda, pp. [257], 258. Lamellibranchiata, pp. [258‑269]. General review of Molluscan Larvæ, pp. [270‑273]. Development of organs, pp. [273‑288].

Chapter X. Polyzoa.

Entoprocta, pp. [292‑297]. Ectoprocta, pp. [297‑305]. Summary and general considerations, pp. [305‑308].

Chapter XI. Brachiopoda.

Development of the layers, pp. [311‑313]. The history of the larva, pp. [313‑317]. Development of organs, p. [317]. General observations on the affinities of the Brachiopoda, pp. [317], 318.

Chapter XII. Chætopoda.

Formation of the germinal layers, pp. [319‑325]. The larval form, pp. [325‑338]. Formation of organs, pp. [338‑342]. Alternations of generations, pp. 342, 343.

Chapter XIII. Discophora.

Formation of layers, pp. [347‑350]. History of larva, pp. [351‑354].

Chapter XIV. Gephyrea.

Gephyrea nuda, pp. [355‑361]. Gephyrea tubicola, pp. [361‑364]. General considerations, p. [364].

Chapter XV.

Chætognatha, pp. [366‑369]. Myzostomea, pp. [369], 370. Gastrotricha, p. [370].

Chapter XVI.

Nematelminthes, pp. [370‑379]. Acanthocephala, pp. [379‑381].

Chapter XVII. Tracheata.

Prototracheata, pp. [382‑387]. Myriapoda, pp. [387‑395]. Insecta, pp. [395‑429]. Embryonic membranes and the formation of the layers, pp. [400‑406]. Formation of the organs, pp. [406‑417]. Special types of larvæ, pp. [417‑419]. Metamorphosis and heterogamy, pp. [420‑429]. Arachnida, pp. [431‑455]. Formation of the layers and general development, pp. [431‑446]. Formation of the organs, pp. [446‑455]. Formation of the layers and embryonic envelopes in the Tracheata, pp. [456‑458].

Chapter XVIII. Crustacea.

History of larval forms, pp. [459‑511]. Branchiopoda, pp. [459‑465]. Malacostraca, pp. [465‑487]. Copepoda, pp. [487‑492]. Cirripedia, pp. [492‑500]. Ostracoda, pp. [500‑502]. Phylogeny of the Crustacea, pp. [502‑511]. The formation of the germinal layers, pp. [511‑521]. Comparative development of organs, pp. [521‑529].

Chapter XIX.

Pœcilopoda, pp. [534‑538]. Pycnogonida, pp. [538], 539. Pentastomida, pp. [539‑541]. Tardigrada, p. [541]. Summary of Arthropodan Development, pp. [541‑543].

Chapter XX. Echinodermata.

Development of the germinal layers, pp. [544‑553]. Development of the larval appendages and metamorphosis, pp. [553‑573]. Summary and general considerations, pp. [573‑576].

Chapter XXI. Enteropneusta. Pp. [579‑583].

Index. Pp. [584‑590].

Appendix.

EMBRYOLOGY.

INTRODUCTION.

Embryology forms a large and important department of Biology. Strictly interpreted according to the meaning of the word, it ought to deal with the growth and structure of organisms during their development within the egg membranes, before they are capable of leading an independent existence. Modern investigations have however shewn that such a limitation of the science would have a purely artificial character, and the term Embryology is now employed to cover the anatomy and physiology of the organism during the whole period included between its first coming into being and its attainment of the adult state.

The subject-matter of the science of Embryology admits of a twofold classification. It may be placed under a series of heads, each dealing either with a special group of organisms, or with a special department of the whole science. If classified in the first of these ways the science will naturally be divided into an Embryology of Plants, and an Embryology of Animals; each of which admits of further subdivision. In the second way the subject falls under two primary heads; viz. Physiological Embryology and Anatomical Embryology.

The present treatise deals only with the Embryology of Animals, and is further confined to those animals known as Metazoa. The science is moreover treated from the morphological or anatomical, rather than from the physiological side.

The marvellous phenomenon of the evolution of a highly complicated living being from a simple undifferentiated germ in which it needs the aid of the most modern microscopical appliances to detect any visible signs of life, has not unnaturally attracted the attention of biologists from the very earliest periods. Before the establishment of the cell theory the origin of the organism from the germ was not known to be an occurrence of the same nature as the growth of the fully formed individual, and Embryological investigations were mixed up with irrelevant speculations on the origin of life[1].

The difficulties of understanding the formation of the individual from the structureless germ led anatomists at one time to accept the view “according to which the embryo preexisted, even though invisible, in the ovum, and the changes which took place during incubation consisted not in a formation of parts, but in a growth, i.e. in an expansion with concomitant changes of the already existing germ.”

Great as is the interest attaching to the simple and isolated life histories of individual organisms, this interest has been increased tenfold by the generalizations of Mr Charles Darwin.

It has long been recognized that the embryos and larvæ of the higher forms of each group pass, in the course of their development, through a series of stages in which they more or less completely resemble the lower forms of the group[2]. This remarkable phenomenon receives its explanation on Mr Darwin’s theory of descent. There are, according to this theory, two guiding, and in a certain sense antagonistic principles which have rendered possible the present order of the organic world. These are known as the laws of heredity and variation. The first of these laws asserts that the characters of an organism at all stages of its existence are reproduced in its descendants at corresponding stages. The second of these laws asserts that offspring never exactly resemble their parents. By the common action of these two principles continuous variation from a parent type becomes a possibility, since every acquired variation has a tendency to be inherited.

The remarkable law of development enunciated above, which has been extended, especially by the researches of Huxley[3] and Kowalevsky, beyond the limits of the more or less artificial groups created by naturalists, to the whole animal kingdom, is a special case of the law of heredity. This law, interpreted in accordance with the theory of descent, asserts that each organism in the course of its individual ontogeny repeats the history of its ancestral development. It may be stated in another way so as to bring out its intimate connection with the laws of inheritance and variation. Each organism reproduces the variations inherited from all its ancestors at successive stages in its individual ontogeny which correspond with those at which the variations appeared in its ancestors. This mode of stating the law shews that it is a necessary consequence of the law of inheritance. The above considerations clearly bring out the fact that Comparative Embryology has important bearings on Phylogeny, or the history of the race or group, which constitutes one of the most important branches of Zoology.

Were it indeed the case that each organism contained in its development a full record of its origin, the problems of Phylogeny would be in a fair way towards solution. As it is, however, the law above enunciated is, like all physical laws, the statement of what would occur without interfering conditions. Such a state of things is not found in nature, but development as it actually occurs is the resultant of a series of influences of which that of heredity is only one. As a consequence of this, the embryological record, as it is usually presented to us, is both imperfect and misleading. It may be compared to an ancient manuscript with many of the sheets lost, others displaced, and with spurious passages interpolated by a later hand. The embryological record is almost always abbreviated in accordance with the tendency of nature (to be explained on the principle of survival of the fittest) to attain her ends by the easiest means. The time and sequence of the development of parts is often modified, and finally, secondary structural features make their appearance to fit the embryo or larva for special conditions of existence. When the life history of a form is fully known, the most difficult part of his task is still before the scientific embryologist. Like the scholar with his manuscript, the embryologist has by a process of careful and critical examination to determine where the gaps are present, to detect the later insertions, and to place in order what has been misplaced.

The aims of Comparative Embryology as restricted in the present work are two-fold: (1) to form a basis for Phylogeny, and (2) to form a basis for Organogeny or the origin and evolution of organs. The justification for employing the results of Comparative Embryology in the solution of the problems in these two departments of science is to be found in the law above enunciated, but the results have to be employed with the qualifications already hinted at; and in both cases a knowledge of Comparative Anatomy is a necessary prelude to their application.

In accordance with the above objects Comparative Embryology may be divided into two departments.

The scientific method employed in both of these departments is that of comparison, and is in fact fundamentally the same as the method of Comparative Anatomy. By this method it becomes possible with greater or less certainty to distinguish the secondary from the primary or ancestral embryonic characters, to determine the relative value to be attached to the results of isolated observations, and generally to construct a science out of the rough mass of collected facts. It moreover enables each observer to know to what points it is important to direct his attention, and so prevents that simple accumulation of disconnected facts which is too apt to clog and hinder the advance of the science it is intended to promote.

In the department of Phylogeny the following are the more important points aimed at.

(1) To test how far Comparative Embryology brings to light ancestral forms common to the whole of the Metazoa. Examples of such forms have been identified by various embryologists in the ovum itself, supposed to represent the unicellular ancestral form of the Metazoa: in the ovum at the close of segmentation regarded as the polycellular Protozoon parent form: in the two-layered gastrula, etc., regarded by Haeckel as the ancestral form of all the Metazoa[4].

(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 Cœlenterata, 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 Cœlenterate 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 amœboid 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.

In order to understand the meaning of conjugation in connection with reproduction, it is important to understand how the two became in the first instance related. For the solution of this question the fact that many Protozoa have the capacity of temporarily or permanently fusing together without an immediate act of reproduction is of great importance. A good example of such fusion is supplied by Actinophrys. We must suppose in fact that the simple coalescence of two or more individuals gives a sufficient amount of extra vigour to their product, to compensate the race for the loss in number of individuals so caused. This extra vigour probably first exhibited itself especially by increased activity in reproduction, till finally the two processes, viz. that of conjugation and that of reproduction, came to be inseparably connected together.

The reproduction of the forms above the Protozoa, which are known as the Metazoa, takes place by two methods, viz. a sexual and an asexual one. The sexual process, which occurs in every known Metazoon[5], consists essentially, as is shewn in the second chapter of this work, in the fusion of two cells budded off from the parent organism, viz. the female cell or ovum, and the male cell or spermatozoon, and of the subsequent division of the compound cell so produced into a number of parts which build themselves up into an organism resembling one of the parents. The sexual process has obviously at first sight a very close resemblance to the process of conjugation. Since it is a question of fundamental importance to determine how sexual reproduction originated, it becomes necessary to examine how far this apparent resemblance is a real one, and how far sexual reproduction can be derived from reproduction following upon conjugation.

In spite of the general similarity between the two processes there is an obvious difficulty in comparing them, in that the result of conjugation is usually the breaking up of the individual formed by the fusion of two other individuals into a number of new organisms, while the result of the fusion which takes place in sexual reproduction is the formation of a single new organism. This difference between the two processes, great as it is, is perhaps apparent rather than real. It must be remembered that a single individual Metazoon is equivalent to a number of Protozoa coalesced to form a single organism in a higher state of aggregation. It results from this that the segmentation of the ovum which follows the sexual act may be compared to the breaking up of the product of conjugation into spores, the difference between the two processes consisting in the fact that in the one case the spores separate each to form an independent organism, while in the other they remain united and give rise to a single compound organism.

If the above considerations are well founded it seems permissible to accept the general view according to which sexual reproduction is derived from conjugation. It is necessary to suppose that, in a colony of Protozoa in the course of becoming a Metazoon, the capacity of reproduction by spores became localized in certain definite cells, and although the formation of spores from these cells may have been possible without previous conjugation, yet that conjugation gradually became established as the rule. The differentiation of primitively similar conjugating cells into male and female cells was probably a very early occurrence, since indications of an analogous differentiation, as has already been mentioned, are found in certain existing Protozoa (Monads, Vorticella, etc.). I have attempted to shew in the second chapter that the breaking up of the cell into spores without previous conjugation is perhaps provided against in the extrusion of the so-called ‘directive body’.

With the differentiation of special germinal cells, to take the place of the whole individual in the act of conjugation, the possibility of each act of conjugation resulting in the production of only a single organism became introduced. Germinal cells can be indefinitely produced, and the reproductive capacity of a single individual is therefore unlimited; while if two whole individuals conjugated and only produced one from the process, the result would be a diminution instead of an increase in the race[6].

It must be admitted that, in the present state of our knowledge, the passage from reproduction by spores following conjugation, to true sexual reproduction, can only be traced in a very speculative manner, and that a further advance in our knowledge may prove that the steps which I have attempted to sketch out are far from representing the true origin of sexual differentiation. The peculiar conjugation and fusion of two individuals to form Diplozoon paradoxum may be alluded to in this connection. This fusion merely results in the attainment of sexual maturity by the two conjugating individuals. It does not appear to me probable that this conjugation is in any way connected with the conjugation of the Protozoa, but the reverse must be borne in mind as a possibility.

It is not easy to decide whether the hermaphrodite or the diœcious state is the primitive one, or in other words whether the two conjugating cells, from which I have supposed the sexual products to originate, were derived in the first instance from one or from two colonies of Protozoa. On purely a priori grounds it seems probable that they were originally formed in one colony, and that their derivation from two colonies or individuals was inaugurated when the spermatozoon became motile. There can be no doubt that the diœcious state is a very early one, and that the majority of existing cases of hermaphroditism are secondary.

The above considerations with reference to the male and female cells appear to indicate that they were primitively homodynamous; a conclusion which is on the whole borne out by the history of their development.

Although the modes of reproduction amongst the Metazoa have been divided into the classes sexual and asexual, there is nevertheless one mode of asexual reproduction which ought to be classified with the sexual rather than with the asexual modes. I mean parthenogenesis, which consists essentially in the development of the ovum into a fresh individual without previous coalescence with the male element. This mode of reproduction, which has a very limited range in the animal kingdom, being confined to the Arthropoda and Rotifera, is undoubtedly secondarily derived from sexual reproduction. The conditions of its occurrence are discussed in the second chapter.

It is remarkable that in certain cases the absence of fertilization causes the production of males (Bees, a Saw-fly, Nematus ventricosus, etc.); more usually it results in the production of females only, and there are very often in the Arthropoda a series of successive generations of females all producing ova which develop parthenogenetically into females; eventually however, usually in direct or indirect connection with a change of food or temperature, or other conditions, ova are formed which give rise without fertilization both to males and females.

The true asexual modes of reproduction amongst the Metazoa consist of fission and gemmation. Gemmation is by far the most widely disseminated of the two. Various as are the methods in which it takes place, it seems nevertheless that cells derived from all the germinal layers, and very frequently from all the important organs of the adult, assist in forming the bud. Into the details of the process, which require in many points a fuller elucidation, it is not my purpose to enter.

Gemmation is a far commoner occurrence amongst the simpler than amongst the more highly organised forms. It appears to have been superadded to the sexual mode of reproduction quite independently in a number of different instances.

While there is no difficulty in understanding how gemmation may have started in such simple types as the Cœlenterata, the manner in which it first originated in certain highly organised forms, as for instance the Ascidians, is somewhat obscure, but it seems probable that it began with the division of the developing germ into two or more embryos, at a very early stage of growth.

Such a division of the germ is, as has been shewn by Kleinenberg, normal in Lumbricus trapezoides[7] and Haeckel has shewn that an artificial division of the germ in the Siphonophora leads to the development of two individuals. It has been pointed out by various naturalists that the production of double monsters is often a phenomenon of the same nature. While it is next to impossible to understand how production of a bud could commence for the first time in the adult of a highly organised form, it is not difficult to form a picture of the steps by which the fission of the germ might eventually lead to the formation of buds in the adult state.

The coexistence of sexual reproduction with normal asexual multiplication, or with parthenogenesis, has led to a remarkable phenomenon in the animal kingdom known as alternations of generations[8].

For the details of the various types of alternations of generations, and their origin, the reader is referred to the body of the work; but a few general remarks on the nature and origin of the process, and on its nomenclature, may conveniently be introduced in this place. The simplest cases are those in which an individual which produces by sexual means gives origin to asexual individuals differently organised to itself, which produce by budding the original sexual form, and so complete a cycle. Instances of this kind are supplied by the Hydrozoa, Annelida and Tunicata. In the case of the Tunicata (Doliolum) two different asexual generations may be interpolated between the sexual generations. In all these cases the origin of the phenomenon is easily understood. It appears, as is most clearly shewn in the case of the Annelida, that the ancestors of the species which now exhibit alternations of generations originally reproduced themselves at the same time both sexually and by budding, though probably the two modes of reproduction did not take place at the same season. Gradually a differentiation became established, by which sexual reproduction was confined to certain individuals, which in most instances did not also reproduce asexually. After the two modes of reproduction became confined to separate individuals, the dissimilarity in habits of life necessitated by their diverse functions caused a difference in their organization; and thus a complete alternation of generations became established. The above is no merely speculative history, since all gradations between complete alternations of generations and simple budding combined with sexual reproduction can be traced in actually existing forms.

The alternation of generations as it is found amongst the Entoparasitic Trematodes and most Cestodes, is to be explained in a slightly different way.

It appears that in these parasitic forms a complicated metamorphosis first arose from the parasite having to accommodate itself to the different hosts it was compelled to inhabit, owing to the liability of its primitive and subsequent hosts to be devoured[9]. A capacity for asexual multiplication—obviously of immense advantage to a parasite—appears to have been acquired in some of the stages of this metamorphosis, and an alternation of generations thus established.

A nearly parallel series to that exhibiting alternations of sexual generations with generations which produce by budding is supplied by the cases where sexual generations alternate with parthenogenetic ones, or in some instances even with larvæ which reproduce sexually or else parthenogenetically.

The best known examples of this form of alternations of generations are found amongst the Insecta[10]. A simple case is that of the Aphides. The ova deposited by impregnated females give rise to forms differently organised to the parents but provided with an ovary[11]. The eggs from the ovary develop parthenogenetically within the oviduct, and so long as there is plenty of food and warmth the generations produced are always parthenogenetic forms. The failure of warmth and nutriment causes the production of true males and females, and so the cycle is completed. We must suppose that the capacity possessed by so many female insects of producing eggs capable of developing without the influence of the male element, has been, so to speak, taken hold of by natural selection, and has led to the production of viviparous parthenogenetic forms, by which, so long as food is abundant, a clear economy in reproduction is effected. The continuance of the species during winter is secured by the production of males and females, the females laying eggs in autumn which are hatched in the spring.

In Chermes there is less modification of the primitive condition in that the parthenogenetic generations lay their eggs like the impregnated females. In the gall-flies (Cynipidæ), there is frequently an alternation of generations of the same kind as in Chermes; there being no viviparous forms. The individuals of the different generations differ from each other to some extent in all these cases.

A second type of alternations of parthenogenetic and sexual generations is exemplified by the cases of Chironomus and Cecidomyia, where the larvæ which develop from the eggs of the fertilized female produce parthenogenetically, by means of true ova, forms which eventually after several generations (Cecidomyia) of larval reproduction give rise to sexual forms. The explanation is here practically the same as in the case of Aphis, and is paralleled in the gemmiparous series by the production of buds in the larval forms of Trematodes, etc. A very similar occurrence takes place in Ascaris nigrovenosa (vide chapter on Nematoidea), except that larval forms, which carry on reproduction and then perish without developing farther, do so by a true sexual process. Thus there is an alternation of generations of adult and larval sexual forms. The Axolotl is an intermittent example of the same phenomenon.

As might be anticipated from the mode in which alternations of generations have become established, incomplete approximations to it are not uncommon. Such approximations are especially found in the Arthropoda, where alternations of sexual and parthenogenetic generations frequently take place, in which the individuals of different generations are similarly organised (Psychidæ, Apus, &c.). Another approximation is afforded by the parthenogenetic winter eggs of Leptodora amongst the Phyllopods, which give rise to Nauplius larvæ, while the young hatched from the summer eggs do not pass through a metamorphosis. Numerous transitional cases are also found amongst the forms in which there is an alternation of sexual and gemmiparous generations.

The whole of the cases to which allusion has been made in this section may be conveniently classed under the term alternations of generations, but the cases of alternation of two sexual generations, and of sexual and parthenogenetic generations, are classified by Leuckart, Claus, etc. as cases of heterogeny, which they oppose to the other form of alternation of generations. If special terms are to be adopted for the two kinds of alternation of generations, it would be perhaps convenient to classify the cases of alternations of sexual and gemmiparous generations under the term metagenesis, and to employ the term heterogamy for the cases of alternation of sexual and parthenogenetic generations.

The term Nurse (German Amme), employed for the asexual generations in metagenesis, may advantageously be dropped altogether.

[1] To this general statement Wolff forms a remarkable exception, for though without any clear knowledge of what we call cells he had very distinct notions on the relations of growth and development.

[2] Von Baer who is often stated to have established the above generalization really maintained a somewhat different view. He held (Ueber Entwickelungsgeschichte d. Thiere, p. 224) that the embryos of higher forms never resembled the adult stages of lower forms but merely the embryos of such forms. Von Baer was mistaken in thus absolutely limiting the generalization, but his statement is much more nearly true than a definite statement of the exact similarity of the embryos of higher forms to the adults of lower ones.

[3] Huxley was the first to shew that the body of the Cœlenterata was formed of two layers, and to identify these with the two primary germinal layers of the Vertebrata.

[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.

[5] Dicyema, if it is a true Metazoon, would seem to form an exception to this rule.

[6] In the vegetable kingdom there are numerous types of Thallophytes, which throw a considerable amount of light on the relation between sexual reproduction and conjugation. Subjoined are a few of the more striking cases. In Pandorina at the time of sexual reproduction the cells which constitute a colony divide each into sixteen, and the products of their division are set free. Pairs of them then conjugate and permanently fuse. After a resting stage the protoplasm is set free from its envelope after division into two or four parts. Each of these then divides into sixteen coherent cells and constitutes a new Pandorina colony. In Œdogonium the fertilization is effected by a spermatozoon fusing with an oosphere (ovum). The fertilized oosphere (oospore) then undergoes segmentation like the ovum of an animal; but the segments, instead of uniting to form a single organism, separate from each other, and each of them gives rise to a fresh individual (swarm-spore) which grows into a perfect Œdogonium. In Coleochæte the impregnation and segmentation take place nearly as in Œdogonium, but the segments remain united together, acquire definite cell walls, and form a single embryo. There is in fact in Coleochæte a true sexual reproduction of the ordinary type. (Vide S. H. Vines “On alternation of generation in the Thallophytes.” Journal of Botany, Nov., 1879.)

[7] The case of Pyrosoma, which might be cited in this connection, is probably secondary.

[8] For an excellent account of this subject, vide Allen Thompson’s article Ovum in Todd’s Cyclopædia. The metamorphosis of the Echinoderms included under this head in Thompson’s article is now known not to be a proper case of alternations of generations.

[9] The appearance of Vertebrata on the globe as the forms which most frequently preyed on Invertebrate forms, and were themselves not so liable to be devoured, has no doubt had a great influence on the metamorphosis of internal parasites, and has amongst other things resulted in these parasites usually reaching their sexual state in a vertebrate host.

[10] For details vide Chapter on Insecta.

[11] The distinction drawn by Huxley between ova and pseudova does not appear to me a convenient one in practice.

CHAPTER I.

THE OVUM AND SPERMATOZOON.

The Ovum.

The complete developmental history of any being constitutes a cycle. It is therefore permissible in treating of this history to begin at any point. As a matter of convenience the ovum appears to be the most suitable point of departure. The question as to the germinal layer from which it is ultimately derived is dealt with in a subsequent part of the work; the present chapter deals with its origin and growth.

General History of the Ovum.

Every young ovum ([fig. 1]) has the character of a simple cell. It is formed of a mass of naked protoplasm (a), containing in its interior a nucleus (b), within which there is a nucleolus (c). The nucleus and nucleolus are usually known as the germinal vesicle and germinal spot.

Fig. 1. Diagram Of the Ovum. (From Gegenbaur.)

a. Granular protoplasm. b. Nucleus (germinal vesicle). c. Nucleolus (germinal spot).

The ovum so constituted is developed either (1) from one cell out of an aggregation or layer of cells all of which have the capacity of becoming ova; or (2) from one of a number of cells segmented off from a polynuclear mass of protoplasm, not divided into separate cells. In both cases the cells which have the capacity of becoming ova may be spoken of as germinal cells, and in the case where the ova are ultimately developed from a polynuclear mass of protoplasm the latter structure may be called a germogen.

In some cases the whole of the germinal cells eventually become ova, but as a rule only a small proportion of them have this fate, the remainder undergoing various changes to be spoken of in the sequel.

Extended investigations have shewn that the distinction between germinal cells which are independent cells from the first, or derived from a germogen in which the nucleated protoplasm is not divided into cells, is an unimportant one; and closely allied forms may differ in this respect. It is moreover probable that a germogen of nucleated protoplasm is less common than is often supposed: it being a matter of great difficulty to determine the structure of the organs usually so described. A germogen is stated to be found in most Platyelminthes, Nematoidea, Discophora, Insecta, and Crustacea.

A more important distinction in the origin of the germinal cells is that afforded by their position. In this respect three groups may be distinguished. (1) The germinal cells may form the lining of a sack or tube, having the form of a syncytium or of an epithelium of separate cells (Platyelminthes, Mollusca, Rotifera, Echinodermata, Nematoidea, Arthropoda). (2) Or they may form a specialized part of the epithelium lining the general body cavity (Chætopoda, Gephyrea, Vertebrata). (3) Or they may form a mass placed between the two elsewhere contiguous primitive germinal layers (Cœlenterata[12]).

Types of transition between the first and second group are not uncommon. Such types, properly belonging to the second group, originate by a special membranous sack continuous with the oviduct being formed round the primitively free patch of germinal cells. Examples of this are afforded by the Discophora, the Teleostei, etc. It is very probable that all the cases which fall under the first heading may have been derived from types which belonged to the second group.

The mode of conversion of the germinal cells into ova is somewhat diverse. Before the change takes place the germinal cells frequently multiply by division. The change itself usually involves a considerable enlargement of the germinal cell, and generally a change in the character of the germinal vesicle, which in most young ova ([fig. 2]) is very large as compared to the body of the ovum. The most complicated history of this kind is that of the ovum of the Craniata. (Vide pp. [56], [57].)

Fig. 2. Ovum Of Carmarina (Geryonia) hastata. (Copied from Haeckel.)

gd. Body of ovum. gv. Germinal vesicle. gm. Germinal spot.

The ovum in its young condition is obviously nothing but a simple cell; and such it remains till the period when it attains maturity.

Nevertheless the changes which it undergoes in the course of its growth are of a very peculiar kind, and, consisting as they do in many instances of the absorption of other cells, have led various biologists to hold that the ovum is a compound structure. It becomes therefore necessary to consider the processes by which the growth and nutrition of the ovum is effected before dealing with the structure of the ovum at all periods of its history.

Fig. 3. Female gonophore of Tubularia Mesembryanthemum. Containing One Large Ovum (ov) and a number of germinal cells (g.c.).

ep. Epiblast (Ectoderm). hy. Hypoblast (Entoderm). ov. Ovum. g.c. Germinal cells.

The ovum is of course nourished like every other cell by the nutritive fluids in which it is surrounded, and special provisions are made for this, in that the ovary is very frequently placed in contiguity with vascular channels. But in addition to such nutrition a further nutrition, the details of which are given in the special part of this chapter, is provided for in the germinal cells which do not become ova.

In the simplest case, as in many Hydrozoa ([fig. 3]), the germinal cells which do become ova are assimilated by the ovum much in the manner of an Amœba.

In other cases the ovum becomes invested by a special layer of cells, which then constitutes what is known as a follicle. The cells which form the follicle are often germinal cells, e.g. Holothuria, Insecta ([fig. 17]), Vertebrata ([fig. 19]). In other cases they seem rather to be adjoining connective-tissue or epithelioid cells, though it is sometimes difficult to draw the line between such cells and germinal cells. Examples of follicles formed of ordinary connective-tissue cells, are supplied by Asterias, Bonellia ([fig. 16]), Cephalopoda ([fig. 14]), etc.

A membrane enclosing the ovum without a lining of cells, as in many Arachnida, vide p. [51], has no true analogy with a follicle and does not deserve the same name.

The function of the follicle cells appears to be, to elaborate nutriment for the growth of the ovum. The follicle cells are not as a rule directly absorbed into the body of the ovum, though in some instances, as in Sepia (vide p. [40]), they are eventually assimilated in this way.

In many cases some of the germinal cells form a follicle, while other germinal cells form a mass within the follicle destined eventually to be used as pabulum. Insects supply the best known examples of this, but Piscicola, Bonellia (?) may also be cited as examples of the same character. In the Craniata (pp. [56‑58]) some of the germinal cells which advance a certain distance on the road towards becoming ova, are eventually used as pabulum, before the formation of the follicle; while other germinal cells form at a later period the follicular epithelium. A peculiar case is that of the Platyelminthes ([fig. 9]), where a kind of follicle is constituted by the cells of a specially differentiated part of the ovary, known as the yolk-gland. The cells of this follicle may either remain distinct, and continue to surround the ovum after its development has commenced, and so be used as food by the embryo; or they may secrete yolk particles, which enter directly into the protoplasm of the ovum.

For further variations in the mode of nutrition the reader is referred to the special part of this chapter. Suffice it to say that none of the known modes of nutrition indicate that the ovum becomes a compound body any more than the fact of an Amœba feeding on another Amœba would imply that the first Amœba ceased thereby to be a unicellular organism.

The constitution of the ovum may be considered under three heads:—

(1) The body of the ovum.
(2) The nucleus or germinal vesicle.
(3) The investing membranes.

The body of the ovum. The essential constituent of the body of the ovum is an active living protoplasm. As a rule there are present certain extraneous matters in addition, which have not the vital properties of protoplasm. The most important of these is known as food-yolk, which appears to be generally composed of an albuminoid matter.

The body of the ovum is at first very small compared with the germinal vesicle, but continually increases as the ovum approaches towards maturity. It is at first comparatively free from food-yolk; but, except in the rare instances where it is almost absent, food-yolk becomes deposited in the form of granules, or highly refracting spheres, by the inherent activity of the protoplasm during the later stages in the ripening of the ovum. In many instances the protoplasm of the ovum assumes a sponge-like or reticulate arrangement, a fluid yolk substance being placed in the meshes of the reticulum. The character of the food-yolk varies greatly. Many of its chief modifications are described below. There is not unfrequently present in the vitellus a peculiar body known as the yolk nucleus, which is very possibly connected with the formation of the food-yolk. It is found in many Arachnida, Myriapoda, Amphibia, etc.[13]

Fig. 4. A. Ovum of Hydra in the amœboid state, with yolk-spherules (pseudocells) and Chlorophyll Granules. (After Kleinenberg.)

gv. Germinal vesicle.

B. Single pseudocell of Hydra.

More important for the subsequent development than the variation in the character of the food-yolk is its amount and distribution. In a large number of forms it is distributed unsymmetrically, the yolk being especially concentrated at one pole of the ovum, the germinal vesicle, surrounded by a special layer of protoplasm comparatively free from food-yolk, being placed at the opposite pole. In the Arthropoda it has in most instances a symmetrical distribution. Further details on this subject are given in connection with the segmentation; the character of which is greatly influenced by the distribution of food-yolk.

The body of the ovum is usually spherical, but during a period in its development it not unfrequently exhibits a very irregular amœboid form, e.g. Hydra ([fig. 4]), Halisarca.

Fig. 5. Unripe ovum of Toxopneustes lividus. (Copied from Hertwig.)

The germinal vesicle. The germinal vesicle exhibits all the essential characters of a nucleus. It has a more or less spherical shape, and is enveloped by a distinct membrane which seems, however, in the living state to be very often of a viscous semi-fluid nature and only to be hardened into a membrane by the action of reagents (Fol). The contents of the germinal vesicle are for the most part fluid, but may be more or less granular. Their most characteristic components are, however, a protoplasmic network and the germinal spots[14]. The protoplasmic network stretches from the germinal spots to the investing membrane, but is especially concentrated round the former. ([Fig. 5].) The germinal spot forms a nearly homogeneous body, with frequently one or more vacuoles. It often occupies an eccentric position within the germinal vesicle, and is usually rendered very conspicuous by its high refrangibility. In many instances it has been shewn to be capable of amœboid movements (Hertwig, Eimer), and is moreover more solid and more strongly tinged by colouring reagents than the remaining constituents of the germinal vesicle.

In many instances there is only one germinal spot, or else one main spot and two or three accessory smaller spots. In other cases, e.g. Osseous Fishes, Echinaster fallax, Eucope polystyla, there are a large number of nearly equal germinal spots which appear to result from the division or endogenous proliferation of the original spot. Sometimes the germinal spots are placed immediately within the membrane of the germinal vesicle (Elasmobranchii and Sagitta). In many Lamellibranchiata, in the earthworm, and in many Chætopoda the components of the germinal spot become separated into two nearly spherical masses ([fig. 12]), which remain in contiguity along a small part of their circumference, and are firmly united together. The smaller of the two parts is more highly refractive than the larger. Hertwig has shewn that the germinal spot is often composed of two constituents as in the above cases, but that the more highly refractive material is generally completely enclosed by the less dense substance. By Fol the germinal spot is stated to be absent in a species of Sagitta, but this must be regarded as doubtful. In young ova the relative size of the germinal vesicle is very considerable. It occupies in the first instance a central position in the ovum, but at maturity is almost always found in close proximity to the surface. Its change of position in a large number of instances is accomplished during the growth of the ovum in the ovary, but in other cases does not take place till the ovum has been laid.

As the ovum attains maturity, important changes take place in the constitution of the germinal vesicle, which are described in the next chapter.

The egg membranes. A certain number of ova when ready to be fertilized are naked cells devoid of any form of protecting covering, but as a rule the ovum is invested by some form of membrane. Such coverings present great variety in their character and origin, and may be conveniently (Ludwig, No. [4]) divided into two great groups, viz. (1) those derived from the protoplasm of the ovum itself or from its follicle, which may be called primary egg membranes; and (2) those formed by the wall of the oviduct or otherwise, such as the egg-shell of a bird, which may be called secondary egg membranes.

Fig. 6. Ovum of Toxopneustes variegatus with the pseudopodia-like processes of the protoplasm penetrating the zona radiata (zr). (After Selenka.)

The primary egg membranes may again be divided into two groups (Ed. van Beneden, No. [1]), viz., (1) those formed by the protoplasm of the ovum, to which the name vitelline membranes will be applied; and (2) those formed by the cells of the follicle, to which the name chorion will be applied.

The secondary egg membranes will be dealt with in connection with the systematic account of the development of the various groups. They coexist as a rule with primary membranes, though in some types (Cephalophorous Mollusca, many Platyelminthes, etc.), they constitute the only protecting coverings of the ovum.

The vitelline membranes are either simple structureless membranes or present numerous radial pores. Membranes with the latter structure are very widely distributed, Echinodermata, Gephyrea, Vertebrata, etc. (Vide figs. [5] and [7].) The function of the pores appears to be a nutritive one. They either serve for the emission of pseudopodia-like processes of the protoplasm of the ovum, as has been very beautifully shown in the case of Toxopneustes by Selenka ([fig. 6]), or they admit (?) processes of the follicular epithelial cells (Vertebrata). Their presence is in fact probably caused by the existence of such processes, which prevent the continuous deposition of the membrane. The term zona radiata will be applied to perforated membranes of this kind. Two vitelline membranes, one perforated and the other homogeneous, may coexist at the same time, e.g. Sipunculida, Vertebrata. ([Fig. 7].)

Fig. 7. Section through a small part of the surface of an ovum of an immature female of Scyllium Canicula.

fe. Follicular epithelium. vt. Vitelline membrane. Zn. Zona radiata. yk. Yolk with protoplasmic network.

The chorion is often ornamented with various processes, etc. It is in many cases doubtful whether a particular membrane is a chorion or a vitelline membrane.

All the membranes which surround the ovum may be provided with a special aperture known as the micropyle. A micropyle is by no means found in the majority of types, and there is no homology between the various apertures so named. Micropyles have two functions, either to assist in the nutrition of the ovum during its development, or (2) to permit the entrance of the spermatozoa. The two functions may in some cases coexist. Micropyles of the first class are developed at the point of attachment of the ovum to the wall of the ovary or to its follicle. Good examples of this kind of micropyle are afforded by the Lamellibranchiata ([fig. 12]), Holothuria, and many Annelida (Polynoe, etc.). The micropyle of the Lamellibranchiata (p. [37]) probably serves also to admit the spermatozoa. The second type of micropyle is found in many Insecta, Teleostei, etc.

General Bibliography of the Ovum.

(1) Ed. van Beneden. “Recherches sur la composition et la signification de l’œuf,” etc. Mém. cour. d. l’Acad. roy. des Sciences de Belgique, Vol. XXXIV. 1870.
(2) R. Leuckart. Artikel “Zeugung,” R. Wagner’s Handwörterbuch d. Physiologie, Vol. IV. 1853.
(3) Fr. Leydig. “Die Dotterfurchung nach ihrem Vorkommen in d. Thierwelt u. n. ihrer Bedeutung.” Oken, Isis, 1848.
(4) Ludwig. “Ueber d. Eibildung im Thierreiche.” Arbeiten a. d. zool.-zoot. Institut Würzburg, Vol. I. 1874[15].
(5) Allen Thomson. Article “Ovum” in Todd’s Cyclopædia of Anatomy and Physiology, Vol. V. 1859.
(6) W. Waldeyer. Eierstock u. Ei. Leipzig, 1870.

Special History of the Ovum in different types.

Cœlenterata.

(7) Ed. van Beneden. “De la distinction originelle d. testicule et de l’ovaire.” Bull. Acad. roy. Belgique, 3^e série, Vol. XXXVII. 1874.
(8) R. and O. Hertwig. Der Organismus d. Medusen. Jena, 1878.
(9) N. Kleinenberg. Hydra. Leipzig, 1872.

Amongst the Cœlenterata the ova are developed in imperfectly specialized organs, which are situated in various parts of the body, for the most part in the space between the epiblast and the hypoblast.

In Hydra the locality where the ova are developed only becomes specialized at the time when an ovum is about to be formed. At one or more points the interstitial cells of the epiblast increase in number and form a protuberance of germinal cells, which may be called the ovary. In this ovary a single ovum is formed by the special growth of one cell. (Kleinenberg, No. [9].) In the free and attached gonophores of Hydrozoa, the ova appear either around the walls of the stomach, or the radial canals, or around other parts of the gastro-vascular canals.

Fig. 8. Ripe Ovum Of Epibulia aurantiaca. The Germinal Vesicle has become invisible without reagents.

Copied from Metschnikoff, “Entwicklung der Siphonophoren.” Zeitschrift f. wiss. Zool., Vol. XXIV. 1874.

p.d. Peripheral layer of denser protoplasm. p.m. Central area consisting of a protoplasmic meshwork.

Their close relations to the gastrovascular canals are probably determined by the greater nutritive facilities thereby afforded. (Hertwig, No. [8].)

In the permanent Medusa forms the ova have similar relations to the gastro-vascular system. Amongst the Actinozoa the ova are usually developed between the epiblast and the hypoblast in the walls of the gastric mesenteries. Amongst the Ctenophora the ova are situated in close relation with the peripheral canals of the gastro-vascular system, which run along the bases of the ciliated bands. There are many examples amongst the Cœlenterata of ova which retain in their mature state the very simple constitution which has been described as characteristic of all young ova; and which are, when laid, absolutely without any trace of a vitelline membrane or chorion. In many other cases both amongst the Medusæ, the Siphonophora, and the Ctenophora, the ripe egg exhibits a distinction into two parts. The outer part is composed of a dense protoplasm, while the interior is composed of a network or more properly a spongework of protoplasm enclosing in its meshes a more fluid substance. ([Fig. 8].)

In some cases the ovum while still retaining the constitution last described becomes invested by a very delicate membrane. Such is the constitution of the ripe ovum of Hippopodius gleba amongst the Siphonophora[16] and of the eggs of Geryonia amongst the permanent Medusæ[17]. The ripe eggs of the Ctenophora usually present a similar structure[18]. After being laid they are found to be invested by a delicate membrane separated by a space filled with fluid from the body of the ovum. The latter is composed of two layers, an outer one of finely granular protoplasm and an inner layer consisting of a protoplasmic spongework containing in its meshes irregular spheres. These latter are stated by Agassiz to be of a fatty nature, and it is probable that in most cases where a protoplasmic network is present, this alone constitutes the active protoplasm and that the substance which fills up its meshes is to be looked on as a form of food-yolk or deutoplasm, though it appears sometimes to have the power of assimilating the firmer yolk particles.

The membrane which invests the ovum of many of the Cœlenterata is probably a vitelline membrane.

The ova of the Hydrozoa take their origin, in most groups at any rate[19], from the deeper layer of the epiblast (interstitial layer of Kleinenberg). The interstitial cells in the ovarian region form primary germinal cells, and by an excess of nutrition certain of them outstrip their fellows and become young ova. Such ova differ from the full-grown ova already described, mainly in the fact that they have a proportionately smaller amount of protoplasm round the germinal vesicle. They grow to a considerable extent at the expense of germinal cells which do not become converted into ova.

The ova of many Cœlenterata undergo changes of a more complicated kind before attaining their full development. Of these ova that of Hydra may be taken as the type. The ovary of Hydra (Kleinenberg, No. [9]) is constituted of angular flattish germinal cells of which no single one can be at first distinguished from the remainder. As growth proceeds one of the cells occupying a central position becomes distinguished from the remaining cells by its greater size, and wedge-like shape. It constitutes the single ovum of the ovary. After it has become prominent it grows rapidly in size, and throws out irregular processes. The germinal vesicle, which for a considerable time remains unaltered, also at length begins to grow; and the sharply defined germinal spot which it contains after reaching a certain size completely vanishes. After the atrophy of the germinal spot, there appears in the middle of the ovum a number of roundish yolk granules.

The shape of the ovum becomes more irregular, and chlorophyll granules, in addition to the yolk granules, make their appearance in it. A fresh germinal spot of circular form also arises in the germinal vesicle. Protoplasmic processes are next thrown out in all directions, giving to the ovum a marvellous amœboid character. ([Fig. 4].) The amœboid form of the ovum serves no doubt to give it a larger surface for nutrition. Coincidently with the assumption of an amœboid form there appear in the ovum a great number of peculiar bodies. They are vesicles with a thick wall bearing a conical projection into the interior which is filled with fluid. ([Fig. 4]B.) These bodies are formed directly from the protoplasm of the ovum, and are to be compared both morphologically and physiologically with the yolk-spherules of such an ovum as that of the Bird. They are called pseudocells by Kleinenberg, and are found with slightly varying characters in many ova of the Hydrozoa.

They first appear as small highly refracting granules; in these a cavity is formed which is at first central but is eventually pushed to one side by the formation of a conical projection from the wall of the vesicle.

After the growth of the ovum is completed the amœboid processes gradually withdraw themselves, and the ovum assumes a spherical form; still however continuing to be invested by the remaining cells of the ovary. It is important to notice that the egg of Hydra retains throughout its whole development the characters of a single cell, and that the pseudocells and other structures which make their appearance in it are not derived from without, and supply not the slightest ground for regarding the ovum as a structure compounded of more than one cell.

The development of the ova of the Tubularidæ, which has been supposed by many investigators to present very special peculiarities, takes place on essentially the same type as that of Hydra, but the germinal vesicle remains permanently very small and difficult to observe. The mode of nutrition of the ovum may be very instructively studied in this type. The process is one of actual feeding, much as an Amœba might feed on other organisms. Adjoining one of the large ova of the ovary there may be seen a number of small germinal cells. ([Fig. 3].) The boundary between these cells and the ovum is indistinct. Just beyond the edge of the ovum the small cells have begun to undergo retrogressive changes; while at a little distance from the ovum they are quite normal (g.c.)[20].

Platyelminthes.

(10) P. Hallez. Contributions à l’Histoire naturelle des Turbellariés. Lille, 1879.
(11) S. Max Schultze. Beiträge z. Naturgeschichte d. Turbellarien. Greifswald, 1851.
(12) C. Th. von Siebold. “Helminthologische Beiträge.” Müller’s Archiv, 1836.
(13) C. Th. von Siebold. Lehrbuch d. vergleich. Anat. d. wirbellosen Thiere. Berlin, 1848.
(14) E. Zeller. “Weitere Beiträge z. Kenntniss d. Polystomen.” Zeit. f. wiss. Zool., Bd. XXVII. 1876.

[Vide also Ed. van Beneden] (No. [1]).

This group, under which I include the Trematodes, Cestodes, Turbellarians and Nemertines, has played an important part in all controversies relating to the nature and composition of the ovum. The peculiarity in the development of the ovum in most members of this group consists in the fact that two organs assist in forming what is usually spoken of as the ovum. One of these is known as the ovary proper, and the other as the vitellarium or yolk-gland. In the sequel the term ovum will be restricted to the product of the first of these organs. In Trematodes the ovary forms an unpaired organ directly continuous with an oviduct into which there open the ducts from paired yolk-glands.

The ovary has a sack-like form and contains in some instances a central lumen (Polystomum integerrimum). At the blind end of the organ is placed the germinal tissue. This part is, according to the accounts of the majority of investigators, formed of a polynuclear mass of protoplasm not divided into distinct cells. Whether it is really formed of undivided protoplasm or not, it is quite certain that a little lower down in the organ distinct cells are found, which have been segmented off from the above mass, and are formed of a large nucleus and nucleolus, surrounded by a delicate layer of protoplasm. These cells are the young ova. They usually assume a more or less angular form from mutual pressure, and, in the cases where the ovary has a lumen, constitute a kind of epithelial lining for the ovarian tube. They become successively larger in passing down the ovary, and, though in most cases naked, are in some instances (Polystomum integerrimum) invested by a delicate vitelline membrane. Eventually the ova pass into the oviduct and become free and at the same time assume a spherical form.

In the oviduct the ovum receives somewhat remarkable investing structures, derived from the organ before spoken of as the yolk-gland. The yolk-gland consists of a number of small vesicles, each provided with a special duct, connected with the main duct of the gland. Each vesicle is lined by an epithelium of cells provided with doubly contoured membranes, and containing nuclei.

As the yolk cells grow older refracting spherules become deposited in their protoplasm, which either completely hide the nucleus, or render it very difficult to see. In the majority of cases the entire cells forming the lining of the vesicles constitute the secretion of the yolk-gland. They invest the ovum, and around them is formed a shell or membrane. In some cases (e.g. Polystomum integerrimum) the yolk cells retain their cellular character and vitality till the embryo is far developed. In other cases they lose their membrane and nucleus shortly after the formation of the egg-shell, and break up into a fluid, holding in suspension a number of yolk granules. A partial disorganisation of the yolk cells can also take place before they surround the ovum; while in some species of Distomum they completely break up before leaving the yolk-gland.

Fig. 9. Generative system of Vortex viridis. (From Gegenbaur, after Max Schultze.)

t. Testis. v.d. Vasa differentia. v.s. Seminal vesicle. p. Penis. u. Uterus. o. Ovary. v. Vagina. g.v. Yolk-glands. r.s. Receptaculum seminis.

There is thus a complete series of gradations between the investment of the ovum by a number of distinct cells, and its investment by a layer of fluid containing yolk-spherules in suspension. In neither the one case nor the other do the investing structures take any share in the direct formation of the embryo from the ovum. Physiologically speaking they play the same part as the white in the fowl’s egg.

The egg-shell, which is usually formed by a secretion of a special shell-gland opening into the oviduct, exhibits one or two peculiarities in the different species of Trematodes. In Amphistomum subclavatum it presents at one extremity a thickened area, which is pierced by a narrow micropyle. In other cases one extremity of the egg-shell is produced into a long process, and sometimes even both extremities are armed in this way. Opercula and other types of armature are also found in different forms.

The mode of development of the ovum in Cestodes is very nearly the same as in Trematodes.

The ovum becomes enveloped in the usual secretion of the yolk-gland; and an egg-shell is always formed by the secretion of a special shell-gland.

Amongst the Turbellarians and Nemertines, there are greater variations in the arrangement of the female generative glands, than in the preceding types. In most of the Rhabdocœla and fresh-water Dendrocœla these organs resemble in their fundamental characters those of the Trematodes and Cestodes. There are present a paired or single ovary and a paired yolk-gland. The general arrangement of the organs is shewn in [fig. 9].

The blind end of the ovaries is usually (Ed. van Beneden, etc.) stated to be formed of a polynuclear protoplasmic basis, but Hallez (No. [10]) has recently insisted that, even at the extreme end of the ovary, the germinal cells are quite distinct, and not confounded together.

With one or two exceptions the yolk cells secreted by the vitellarium retain their vitality till they are swallowed by the embryo, after the development of its mouth. The few not so swallowed become disintegrated. They are granular nucleated cells, and, as was first shewn by von Siebold, are remarkable for exhibiting spontaneous amœboid movements.

Very important light on the nature of the vitellarium is afforded by the structure of the generative organs in Prorhyncus and Macrostomum.

In Prorhyncus there is no separate vitellarium, but the lower part of the ovarian tube functionally and morphologically replaces it. The ovum becomes surrounded by yolk cells, which according to Hallez (No. [10]) retain their vitality for a long time. According to Ed. van Beneden yolk-spherules are formed in the protoplasm of the ovum itself, in addition to and independently of the surrounding yolk cells. In Convoluta paradoxa a special vitellarium is stated to be absent; though a deposit of yolk is formed round the ovum (Claparède).

In Macrostomum again the yolk-glands are at most represented by a lower specialized part of the ovarian tube. The ova in passing down become filled with yolk-spherules. According to Ed. van Beneden these spherules are formed in the protoplasm of the ovum itself; but this is explicitly denied by Hallez, who finds that they are formed from the lining cells of the ovarian tube, which, instead of retaining their vitality as in Prorhyncus, break up and form a granular mass which is absorbed by the protoplasm of the ovum.

In Prostomum caledonicum (Ed. van Beneden) the generative organs are formed on the same plan as in other Rhabdocœla, but the cells which form the yolk-gland give rise to yolk particles which enter the ovum, instead of to a layer of yolk cells surrounding the ovum.

Amongst the marine dendrocœlous Turbellarians the ova are formed in separate sacks widely distributed in the parenchyma of the body between the alimentary diverticula. In these the ova undergo their complete development, without the intervention of yolk-glands.

The ovaries of the Nemertines more nearly resemble those of the marine Dendrocœla than those of the Rhabdocœla. They consist of a series of sacks situated on the two sides of the body between the prolongations of the digestive canal. The eggs are developed in these sacks in a perfectly normal manner, and in many cases become filled with yolk-spherules which arise as differentiations of the protoplasm of the ovum. The protecting membranes of the ova have not been accurately studied. In some cases[21] two membranes are present, an internal and an external. The former, immediately investing the vitellus, is very delicate: the external one is thicker and hyaline.

The constitution of the female generative organs of the Trematodes was first clearly ascertained by von Siebold (No. [12]). He originally, though not very confidently, propounded the view that the germinal vesicles alone were formed in the ovary and that the protoplasm of the ovum was supplied by the yolk-gland. This view has long been abandoned, and von Siebold (No. [13]) himself was the first to recognize that true ova with a protoplasmic body containing a germinal vesicle and germinal spot were formed in the ovary. The Trematodes have however not ceased to play an important part in forming the current views upon the development of ova, and have quite recently served Ed. van Beneden as his type in exposing his general view upon this subject.

His view consists fundamentally in regarding the secretion of the yolk-glands, which in most cases merely invests the ovum, as homologous with the yolk-spherules which fill the protoplasm of many eggs; and he considers the part of the ovary where in most forms the ova receive their supply of yolk particles, as equivalent to the vitellarium of the Platyelminthes. He further appears to regard the primitive state as that exemplified in Trematodes, Cestodes, etc., and holds that the ovarian types characteristic of other forms are secondarily derived from this, by the coalescence of the primitively distinct vitellarium with the ovary proper.

This appears to me a case of putting the cart before the horse. To my mind the vitellarium is to be regarded, as has already been suggested by Gegenbaur, Hallez, etc. as a special differentiation of the primitively simple ovarian tube, and the instances Of Macrostomum and Prorhyncus just cited appear to me to indicate some of the steps in this differentiation. In Macrostomum the cells of the lower part of the oviduct simply supply a kind of nutriment to the ovum in the form of granular yolk particles, while in Prorhyncus the yolk cells of the lower part of the ovarian tube form a complete investment of independent cells for the ovum. If this lower part of the ovarian tube were to grow out as a special diverticulum we should have produced a normal vitellarium. But even with the above modification the theory of van Beneden appears to me not completely satisfactory. The view that the yolk-spherules are of the same nature as the yolk cells is mainly supported by the case of Prostomum caledonicum, where the vitellarium produces the yolk particles which fill the ovum. The cases Of Prorhyncus and Macrostomum give a different complexion to that of Prostomum caledonicum. From the first of these especially it appears that, even when normal yolk cells surround the ovum, yolk particles can be deposited independently in the protoplasm of the ovum.

The most probable view of the nature of the vitellarium is that of Gegenbaur, Hallez, etc., according to which it is to be regarded as a specially modified part of the ovarian tube. On this view the nature and function of the yolk cells admit of a fairly simple explanation. They are to be regarded as primary germinal cells like those in the ovaries of Hydra, Tubularia, etc., which do not become converted into ova. Like these cells they may in some instances, Macrostomum, Prostomum, etc., serve directly in the nutrition of the ovum. In other cases they retain their independence and serve for the late nutrition of the embryo. In both instances they retain the faculty, normally possessed by ova, of forming yolk particles in their protoplasm.

Echinodermata.

(15) C. K. Hoffmann. “Zur Anatomie d. Echiniden u. Spatangen.” Niederländisch. Archiv f. Zoologie, Vol. I. 1871.
(16) C. K. Hoffmann. “Zur Anatomie d. Asteriden.” Niederländisch. Archiv f. Zoologie, Vol. II. 1873.
(17) H. Ludwig. “Beiträge zur Anat. d. Crinoiden.” Zeit. f. wiss. Zool., Vol. XXVIII. 1877.
(18) Joh. Müller. “Ueber d. Canal in d. Eiern d. Holothurien.” Müller’s Archiv, 1854.
(19) C. Semper. Holothurien. Leipzig, 1868.
(20) E. Selenka. Befruchtung d. Eies v. Toxopneustes variegatus, 1878.

[Vide also Ludwig (No. [4]), etc.]

The eggs of the Echinodermata present in their development certain points of interest.

The ovaries themselves are usually surrounded by a special vascular dilatation. In the Asteroidea, the Echinoidea, and the Holothuroidea the organs have the form of sacks; specially surrounded in the two former groups, and probably the latter, by a vascular sinus formed as a dilatation of one of the generative vessels. In the Crinoids they have the form of a hollow rachis completely surrounded by a blood-vessel. ([Fig. 11], b.) The proximity of the ovaries (generative organs) to the vascular system in these forms has clearly the same physiological significance as the proximity of the ovaries (generative organs) to the radial vessels in the Cœlenterata.

In the Asteroidea, the Echinoidea and the Holothuroidea the ovaries have the form of sacks lined by an epithelium of germinal cells, and the ova are formed by the enlargement of these cells, which, when they have reached a certain size, become detached from the walls, and fall into the cavity of the ovarian sack. In Toxopneustes (Selenka) and very probably in other forms only a few of the epithelial cells undergo conversion into ova: the remainder undergo repeated division, and, as in so many other cases, are eventually employed in the nutrition of the true ova. In the nearly ripe ova of Asterias Fol has described a flattened follicular epithelium the origin of which is unknown.

Fig. 10. Ovum of Toxopneustes variegatus with the pseudopodia-like projections of the protoplasm penetrating the zona radiata (zr). (After Selenka.)

In Holothuria (Semper) a further differentiation of the germinal cells, not destined to become ova, takes place. They surround the enlarged cell which forms the true ovum, for which they constitute a kind of follicular capsule. This capsule is attached by a stalk to the walls of the ovary, and the ovum lies freely in it except for an area nearly opposite its (the capsule’s) point of attachment, where the ovum adheres to the wall of the capsule. Subsequently the follicle cells which form the capsule fuse together, and form a definite membrane in which only the nuclei remain distinct. Within the membranous capsule there is formed for the ovum an albuminous zona radiata. At the point where the ovum is attached to its capsule this membrane cannot be developed, and therefore remains incomplete. The perforation so formed, becomes the micropyle of the Holothurian egg, which was first discovered by Joh. Müller. The albuminous membrane just described for Holothurians is also found in Asteroids ([fig. 5]) and Echinoids. In these groups there is no proper micropyle, though in Ophiothrix a nutritive passage perforates the membrane at the attachment of the ovum before the period when the ovum becomes free (Ludwig). The formation of the zona radiata has been studied by Selenka. It is secreted by the protoplasm of the ovum, and has a gelatinous consistency, and after it is formed the peripheral layer of the protoplasm of the ovum sends out through it pseudopodia-like processes to absorb nutriment from without. These processes are at first large and irregular, but soon become finer and finer ([fig. 10]), and acquire a regular radiating arrangement. They are withdrawn when the ovum is ripe, but they nevertheless give rise to the finely radiated appearance of the membrane, the radii being in reality delicate pores.

Fig. 11. Transverse section through the pinna of a sexually mature Comatula. (From Gegenbaur, after Ludwig.)

p. Tentacle. g. Lumen of genital rachis. w. Water-vascular vessel. n. Nerve cord. b. Blood-vessel on nerve cord and round genital rachis. cg. Genital canal. cd. Dorsal section of the body cavity. cv. Ventral section of body cavity.

In the Crinoids the generative rachis consists of a tube, the epithelium of which is formed of the primary germinal cells. ([Fig. 11].) While some of these cells enlarge and become ova, the remainder supply the elements for a follicular epithelium, which is established round the ova, exactly as in Holothurians.

Mollusca.

Lamellibranchiata.

(21) H. Lacaze-Duthiers. “Organes génitaux des Acéphales Lamellibranches.” Ann. Sci. Nat., 4^me série, Vol. II. 1854.
(22) W. Flemming. “Ueb. d. er. Entwick. am Ei d. Teichmuschel.” Archiv f. mikr. Anat., Vol. X. 1874.
(23) W. Flemming. “Studien üb. d. Entwick. d. Najaden.” Sitz. d. k. Akad. Wiss. Wien, Vol. LXXI. 1875.
(24) Th. von Hessling. “Einige Bemerkungen, etc.” Zeit. f. wiss. Zool., Bd. V. 1854.
(25) H. von Jhering. “Zur Kenntniss d. Eibildung bei d. Muscheln.” Zeit. f. wiss. Zool., Vol. XXIX. 1877.
(26) Keber. De Introitu Spermatozoorum in ovula, etc. Königsberg, 1853.
(27) Fr. Leydig. “Kleinere Mittheilung etc.” Müller’s Archiv, 1854.

Gasteropoda.

(28) C. Semper. “Beiträge z. Anat. u. Physiol. d. Pulmonaten.” Zeit. f. wiss. Zool., Vol. VIII. 1857.
(29) H. Eisig. “Beiträge z. Anat. u. Entwick. d. Pulmonaten.” Zeit. f. wiss. Zool., Vol. XIX. 1869.
(30) Fr. Leydig. “Ueb. Paludina vivipara.” Zeit. f. wiss. Zool., Vol. II. 1850.

Cephalopoda.

(31) Al. Kölliker. Entwicklungsgeschichte d. Cephalopoden. Zurich, 1844.
(32) E. R. Lankester. “On the developmental History of the Mollusca.” Phil. Trans., 1875.

Lamellibranchiata.

The ova of the Lamellibranchiata present several points of interest. They are developed in pouches of the ovary which are lined by a flattened germinal epithelium, or sometimes (?) a syncytium. Some of the cells of this epithelium enlarge and become ova, but remain attached to the walls of their pouches by protoplasmic stalks. Round the ovum there appears in some forms (Anodon, Unio) a delicate vitelline membrane, which is incomplete at the protoplasmic stalk, and is therefore perforated by an aperture which forms the micropyle. ([Fig. 12].) As the ovum becomes ripe a large space filled with albuminous fluid becomes established between the ovum and its membrane, but the ovum remains attached to the membrane at the micropyle. In Scrobicularia (von Jhering, No. [25]) the membrane round the ovum appears from the first as an albuminous layer, the outermost stratum of which becomes subsequently hardened as the vitelline membrane. In this form also the protoplasmic stalk becomes, in pouches largely filled with ova, extremely long. The ova become eventually detached by the stalk rupturing, and the portion of it which remains attached to the vitelline membrane falling off. The function of the stalk and of the micropyle during the development of the ovum is undoubtedly a nutritive one.

In Anodon and Unio yolk granules similar to those deposited in the protoplasm of the ovum are also found in the epithelial cells of the ovarian pouches (Flemming, [22]), and there can be but little doubt that they are directly transported from these cells into the ovum. These cells would seem therefore to play much the same part as the yolk-glands of some Turbellarians (Prostomum caledonicum). In Scrobicularia yolk granules are not found in the epithelium of the pouches, but are contained in the dilated disc by which the ovum is attached to the wall of its pouch, as well as in the ovum itself.

Fig. 12. Medium-sized ovum of Anodonta complanata. (After Flemming.)

mp. micropyle. gs. germinal spot.

On the ovum becoming detached the micropyle still remains as an aperture, which probably has the function of admitting the spermatozoa.

The shape and form of the micropyle vary greatly. In Anodon and Unio it is a projecting trumpet-shaped structure, which after fertilization becomes shortened and reduced to a mere aperture which is finally stopped up. ([Fig. 12].)

In other forms it is simply a perforation in the vitelline membrane which is sometimes very large. In a species of Arca, which I had an opportunity of observing at Valparaizo, it was equal to nearly the circumference of the ovum.

The eggs of the Lamellibranchiata are not only remarkable in the possession of a micropyle, but in certain peculiarities of the yolk and of the germinal vesicle.

In the fresh-water mussels there is usually found in young and medium-sized ova a peculiar lens-shaped body—Keber’s corpuscle—which is placed immediately internal to the micropyle. It is probably in some way connected with the nutrition of the ovum, though the fact that it is not always present shews that it cannot be of great importance.

A dark body found by von Jhering in the neighbourhood of the germinal vesicle in the ripe ovum of Scrobicularia is probably of a similar nature to Keber’s corpuscle. Both bodies may be placed in the same category as the so-called yolk nucleus of the spider’s and frog’s ova.

In all except the youngest ova of Anodon and Unio the germinal spot is composed of two nearly complete spheres united together for a small part of their circumference. ([Fig. 12], gs.) The smaller of these has a higher refractive index than the larger, and often contains a vacuole: the two parts together appear to be the separated components (though not by simple division) of the primitive nucleolus. A nucleolus of this character is not universal amongst Lamellibranchiata, but a similar separation of the constituents of the germinal spot has been found by Flemming in Tichogonia, in which however the more highly refracting body envelopes part of the less highly refracting body in a cap-like fashion.