The title of the Nobel Lecture of George Palade (1 August, p. 347) should have been "Intracellular aspects of the process of protein secretion."

https://science.sciencemag.org/content/sci/189/4200/347.full.pdf

Intracellular aspects of the process of protein synthesis

Oocyte growth and development is an important issue in fish and fisheries biology. This paper reviews the information available on oocyte growth patterns and the rates and dynamics of oocyte growth in teleosts. In synchronous spawners, the weight of the gonad may represent as much as 40% of the overall body weight of the fish. In asynchronous spawners, the weight of the mature ovary is considerably less than in synchronous ovulators, but the ovary shows a more regular periodicity and may grow repeatedly many times during the breeding season. There is a huge variability in egg size in teleosts, with the largest known measuring up to 8 cm in diameter. Within the limits of variance set by genetic constraints, egg size may vary between populations of the same species. Oocytes in all teleosts undergo the same basic pattern of growth: oogenesis, primary oocyte growth, cortical alveolus stage, vitellogenesis, maturation and ovulation. The mechanisms that control oocyte growth are addressed in this review, albeit that the available information, as in all other vertebrates, is very limited. The main hormones that have been shown to affect ovarian growth are gonadotrophin, thyroid hormones, growth hormone, insulin and insulin-like growth factors. An overview of the determinants of fecundity, with particular reference to oocyte recruitment and atresia, is the focus of the second part of the paper. Genetics and nutrition have major effects on fecundity, and studies so far suggest that the determinants of fecundity usually operate during the early part of gametogenesis. The role of atresia in determining fecundity is less clear. The final part of this review highlights some areas of study that are priorities for research on ovarian development in fish.

Oocyte growth and development in teleosts

The spermatozoa of most invertebrates (e.g., sea urchins) and nonmammalian vertebrates (e.g., fishes and amphibians) have full capacity to fertilize eggs upon leaving the testis. Testicular spermatozoa of mammals, on the other hand, do not possess the ability to do so. Their fertilizing capacity develops as they pass through the epididymis (Young, 1931; Nishikawa and Waide, 1952; Blandau and Rumery, 1964; Bedford, 1966; Orgebin-Crist, 1967). This process, apparently “unique” to mammals, is referred to as the epididymal maturation of spermatozoa. Even after their maturation, however, spermatozoa require an additional phase of maturation or capacitation within the female genital tract before they are able to fertilize eggs (Austin, 1951, 1952; Chang, 1951a). Thus, epididymal maturation and capacitation are two extra steps that mammaliam spermatozoa must take before they can effect fertilization. In this chapter, I will discuss how mammalian spermatozoa prepare themselves for fertilization and how the spermatozoa and eggs interact during fertilization. The process and mechanisms of sperm transport in the female genital tract will not be dealt with extensively here. Readers are referred to Bishop (1961, 1969), Blandau (1969), Bedford (1970b, 1972b), Thibault (1972, 1973a), Zamboni (1972), Blandau and Gaddum-Rosse (1974), Hafez and Thibault (1975), Overstreet and Katz (1977), Overstreet and Cooper (1978, 1979b), Overstreet et al. (1978), Shalgi and Kraicer(1978), Cooper et al. (1979) and Hunter (1975, 1980). The rejection or elimination of extra spermatozoa by the fertilized egg, one of the most fascinating events in fertilization, will not be discussed here. Instead, readers are referred to the chapter by Dr. Wolf in this volume. The physiology of egg activation has been described and discussed to some extent by Gwatkin (1977) and Yanagimachi (1978a). There are numerous reviews dealing with general aspects of mammalian fertilization. The following are recommended to aid in grasping the outline of mammalian fertilization: Austin and Bishop (1957), Austin and Walton (1960), Austin (1961, 1968), Blandau (1961), Piko (1969), Thibault (1969), Bedford (1970a,b, 1972b), Gwatkin (1976, 1977), Yanagimachi (1977, 1978a), Bedford and Cooper (1978) and Hunter (1980).

Mechanisms of Fertilization in Mammals

THE author of this book ranks as the foremost American worker on insect morphology. His contributions on the subject are notable for their clarity and originality of thought, and the appearance of a volume, embodying his ideas in comprehensive form, is sure of a hearty welcome. In its preparation, Mr. Snodgrass has incorporated the results of much first-hand study with those of many recent investigators in the same field. He has produced an outstanding book wherein knowledge of facts is combined with that of function and, at the same time, theoretical conceptions of the origins and relationships of organs and parts are not overlooked. Principles of Insect Morphology
 By R. E. Snodgrass. (McGraw-Hill Publications in the Zoological Sciences.) Pp. ix + 667. (New York and London: McGraw-Hill Book Co., Inc., 1935.) 36s. net.

Principles of Insect Morphology

A comparison is made between certain fibrous and regularly twisted biological materials and certain ordered liquids commonly called 'esteric liquid crystals' Three examples of twisted arrangements (Arthropod cuticle, Ascidian tunica, Dinoflagellate chromosomes) and typical textures of cholesteric mesophases are studied for their optical properties and their defects. These materials are strongly different. Very long polymer chains occur in the organic matrix of skeletal tissues or in chromosomes. On the contrary, in a cholesteric liquid crystal, the molecules are free to move one along the other. However the geometry of such systems is similar. The objections to the twisted model are reviewed and evidence is presented to support a generalized twisted model. A list of the known biological cholesteric analogues is given.

Twisted fibrous arrangements in biological materials and cholesteric mesophases.

A morphological basis for transcellular potassium transport in the midgut of the mature fifth instar larvae of Hyalophora cecropia has been established through studies with the light and electron microscopes. The single-layered epithelium consists of two distinct cell types, the columnar cell and the goblet cell. No regenerative cells are present. Both columnar and goblet cells rest on a well developed basement lamina. The basal portion of the columnar cell is incompletely divided into compartments by deep infoldings of the plasma membrane, whereas the apical end consists of numerous cytoplasmic projections, each of which is covered with a fine fuzzy or filamentous material. The cytoplasm of this cell contains large amounts of rough endoplasmic reticulum, microtubules, and mitochondria. In the basal region of the cell the mitochondria are oriented parallel to the long axes of the folded plasma-lemma, but in the intermediate and apical portions they are randomly scattered within the cytoplasmic matrix. Compared to the columnar cell, the goblet cell has relatively little endoplasmic reticulum. On the other hand, the plications of the plasma membrane of the goblet cell greatly exceed those of the columnar cell. One can distinguish at least four characteristic types of folding: (a) basal podocytelike extensions, (b) lateral evaginations, (c) apical microvilli, and (d) specialized cytoplasmic projections which line the goblet chamber. Apically, the projections are large and branch to form villus-like units, whereas in the major portion of the cavity each projection appears to contain an elongate mitochondrion. Junctional complexes of similar kind and position appear between neighboring columnar cells and between adjacent columnar and goblet cells as follows: a zonula adherens is found near the luminal surface and is followed by one or more zonulae occludentes. The morphological data obtained in this study and the physiological information on ion transport through the midgut epithelium have encouraged us to suggest that the goblet cell may be the principal unit of active potassium transport from the hemolymph to the lumen of the midgut. We have postulated that ion accumulation by mitochondria in close association with plicated plasma membranes may play a role in the active movement of potassium across the midgut.

https://rupress.org/jcb/article-pdf/31/1/107/1383597/107.pdf

ACTIVE TRANSPORT BY THE CECROPIA MIDGUT II. Fine Structure of the Midgut Epithelium

This paper presents morphological evidence on the origin of cortical granules in the oocytes of Arbacia punctulata and other echinoderms. During oocyte differentiation, those Golgi complexes associated with the production of cortical granules are composed of numerous saccules with companion vesicles. Each element of the Golgi complex contains a rather dense homogeneous substance. The vesicular component of the Golgi complex is thought to be derived from the saccular member by a pinching-off process. The pinched-off vesicles are viewed as containers of the precursor(s) of the cortical granules. In time, they coalesce and form a mature cortical granule whose content is bounded by a unit membrane. Thus, it is asserted that the Golgi complex is involved in both the synthesis and concentration of precursors utilized in the construction of the cortical granule. Immediately after the egg is activated by the sperm the primary envelope becomes detached from the oolemma, thereby forming what we have called the activation calyx (see Discussion). Subsequent to the elaboration of the activation calyx, the contents of cortical granules are released (cortical reaction) into the perivitelline space. The discharge of the constituents of a cortical granule is accomplished by the union of its encompassing unit membrane, in several places, with the oolemma.

/pdf/oocyte-differentiation-in-the-sea-urchin-arbacia-punctulata-f08uw7mi12.pdf

Oocyte differentiation in the sea urchin, arbacia punctulata, with particular reference to the origin of cortical granules and their participation in the cortical reaction

Fertilization events following coalescence of the gamete plasma membranes and culminating in the formation of the zygote nucleus were investigated by light and electron microscopy in the sea urchin, Arbacia punctulata. Shortly after the spermatozoon passes through the fertilization cone, it rotates approximately 180° and comes to rest lateral to its point of entrance. Concomitantly, the nonperforated nuclear envelope of the sperm nucleus undergoes degeneration followed by dispersal of the sperm chromatin and development of the pronuclear envelope. During this reorganization of the sperm nucleus, the sperm aster is formed. The latter is composed of ooplasmic lamellar structures and fasciles of microtubules. The male pronucleus, sperm mitochondrion, and flagellum accompany the sperm aster during its migration. As the pronuclei encounter one another, the surface of the female pronucleus proximal to the advancing male pronucleus becomes highly convoluted. Subsequently, the formation of the zygote nucleus commences with the fusion of the outer and the inner membranes of the pronuclear envelopes, thereby producing a small internuclear bridge and one continuous, perforated zygote nuclear envelope.

/pdf/the-fine-structure-of-pronuclear-development-and-fusion-in-4y7p9nao7p.pdf

The fine structure of pronuclear development and fusion in the sea urchin, arbacia punctulata

The ovary of the roach Periplaneta americana has been studied by techniques of light and electron microscopy. Each ovariole (panoistic type) contains a linear array of oocytes in varying stages of development. Newly formed oocytes become encased by a layer of follicle cells and begin pinocytosis. All subsequent growth stages of the oocytes are dependent, in part, on this phenomenon. All of the pinocytotic caveolae show an unique surface modification; i.e., on their internal surface they have an amorphous or filamentous substance and their external surface is studded with many fine radially oriented spike-like projections. The pinosomes of early oocytes do not contain a demonstrable internal structure; they are thought to contain nutritive substances for the developing oocytes rather than yolk precursors. When the oocyte enters its last stage of growth, characterized by yolk deposition, the caveolae become filled with a dense material which is thought to be the precursors of yolk. Hence the conclusion is drawn that yolk formation is independent of any cytoplasmic organelle system of the oocyte and that the precursors of this deutoplasmic substance are manufactured outside the ovary and are internalized by the process of pinocytosis. Under the phase-contrast microscope the nucleoli of early oocytes are large irregular masses and show the phenomenon of nucleolar emission (fragmentation). These "emissions" become randomly dispersed in the nucleoplasm and some of them come to be intimately associated with the fenestrated nuclear envelope. After this process ceases, the main nucleolar mass becomes vacuolated. Electron micrographs suggest that the constituent particles of the nucleolar emissions migrate from the nucleus through patent pores of the nuclear envelope.

https://rupress.org/jcb/article-pdf/20/1/131/1367904/131.pdf

Oocyte differentiation and vitellogenesis in the roach periplaneta americana.

The differentiation of the primary envelope of oocytes of the seahorse (Hippocampus erectus) and the pipefish (Syngnathus fuscus) has been investigated by techniques of light- and electron microscopy. The developing oocytes have been divided into four stages according to size. Oogonia are designated as stage I; stages II and III are oocytes; stage IV represents mature eggs. The primary envelope which is produced by the oocyte is initially a tripartite structure; for convenience of description, the portions are referred to as zones 1, 2, and 3, respectively. Zone 1 first appears as a homogeneous substance at approximately the middle of the long axis of each microvillus. Zone 2 is immediately beneath zone 1 and consists of an extremely electron-opaque granular component. Zone 3 is subjacent to zone 2; it is the largest and most complex of the three. Zone 3 consists of an amorphous material organized in a reticular-like network. Staining procedures indicate that the envelope is composed of a glycoprotein. Just before the oocyte matures there is a structural alteration in zones 2 and 3. Zone 2 becomes a compact, dense layer and zone 3 becomes multilaminate. Subsequent to these changes, zone 1 degenerates. The classification of egg envelopes is discussed, and comparisons are made between the primary envelope of the teleosts investigated and the primary envelopes of other species.

/pdf/the-formation-of-the-primary-envelope-during-oocyte-2hut0gam78.pdf

Everett Anderson

Papers

ACTIVE TRANSPORT BY THE CECROPIA MIDGUT II. Fine Structure of the Midgut Epithelium

Oocyte differentiation in the sea urchin, arbacia punctulata, with particular reference to the origin of cortical granules and their participation in the cortical reaction

The fine structure of pronuclear development and fusion in the sea urchin, arbacia punctulata

Oocyte differentiation and vitellogenesis in the roach periplaneta americana.

The formation of the primary envelope during oocyte differentiation in teleosts.