The Fine Structure of the Epithelial Cells of the Mouse Prostate : II. Ventral Lobe Epithelium

The fine structure of the epithelial cells of one component of the prostatic complex of the mouse—the ventral lobe—has been investigated by electron microscopy. This organ is composed of small tubules, lined by tall simple cuboidal epithelium, surrounded by smooth muscle and connective tissue. Electron micrographs of the epithelial cells of the ventral lobe show these to be limited by a cell membrane, which appears as a continuous dense line. The nucleus occupies the basal portion of the cell and the nuclear envelope consists of two membranes. The cytoplasmic matrix is of moderately low density. The endoplasmic reticulum consists of elongated, circular, and oval profiles representing the cavities of this system bounded by rough surfaced membranes. The Golgi apparatus appears localized in a region between the apical border and the nucleus, and is composed of the usual elements found in secretory cells (3, 9). At the base of the cells, a basement membrane is visible in close contact with the outer aspect of the cell membrane. A space of varying width, which seems to be occupied by connective tissue, separates the epithelial cells from the surrounding smooth muscle fibers and the blood vessels. Bodies with the appearance of portions of the cytoplasm, mitochondria, or profiles of the endoplasmic reticulum can be seen in the lumina of the acini and on the bases of these pictures and others of the apical region the mechanism of secretion by these cells is discussed. The fine structural organization of these cells is compared with that of another component of the mouse prostate—the coagulating gland.     

An analysis of the ultrastructure of epithelial cells of the rat proximal tubule at early stages of penicillin elimination

A study was made of the ultrastructure of epithelial cells of the rat proximal tubules nephrons 25-30 seconds, and 15 and 30 minutes following penicillin injection. The morphological changes in the transporting cells that were observed involved the dilation of the inner cavities of canals and cisterns of the endoplasmic reticulum. The number of elements of the smooth endoplasmic reticulum concentrated on the periphery of the cell, near the basolateral border, was seen to increase. Microbodies appeared enlarged with the appearance of the nucleolus; the electron density of the cytoplasmic ground substance fell down. The space between cells augmented, and the interstitial space between infoldings of the basal plasmalemma was much broadened. Similar morphological changes but different in the intensity were found at either stage examined. The maximum changes were established 25-30 seconds following penicillin injection.     

NEW CYTOPLASMIC COMPONENTS IN ARTERIAL ENDOTHELIA

A hitherto unknown rod-shaped cytoplasmic component which consists of a bundle of fine tubules, enveloped by a tightly fitted membrane, was regularly found in endothelial cells of small arteries in various organs in rat and man. It is about 0.1 µ thick, measures up to 3 µ in length, and contains several small tubules, ~150 A thick, embedded in a dense matrix, and disposed parallel to the long axis of the rod. In some of these cells, the cisternae of the endoplasmic reticulum are greatly distended by the accumulation of a dense, finely granular material. The nature and significance of these cytoplasmic components are yet unknown.     

The relationship between the morphology of cell organelles and kinetics of the secretory process in male sex accessory glands of mice

Summary Two male sex accessory glands of the mouse, seminal vesicle and coagulating gland, were compared with the aim of relating differences in the morphology of organelles to the kinetics of the secretory process. The epithelial cells of the two glands were assessed by morphometric analysis, cytochemical staining, and electron-microscopic autoradiography after administration of a labeled amino acid. The rough endoplasmic reticulum of the seminal vesicle comprised narrow parallel cisternae, while that of the coagulating gland was greatly distended and occupied a much larger percentage of the cytoplasmic volume. Radioactively labeled products were secreted much more rapidly in the seminal vesicle than in the coagulating gland. The primary point of difference in kinetics of intracellular transport between the two glands was in exit of material from the rough endoplasmic reticulum. The more rapid drainage of the rough endoplasmic reticulum may be related to its relatively greater membrane surface density and lesser internal volume. In contrast, similarities in size and cytochemical staining in the Golgi apparatus of the two glands were accompanied by similar kinetics of intracellular transport of secretory protein through this organelle.     

The fine structure of the light organ of the New Zealand glow-worm Arachnocampa luminosa (Diptera: Mycetophilidae).

The swollen distal tips of the Malpighian tubules of the glow-worm Arachnocampa luminosa constitute the light organ. The ventral and lateral surfaces are covered by a tracheal ‘reflector’ and the nervous supply to the light organ comes from the ganglion in the penultimate segment. Fine nerve terminals, axons, and glial cells can be seen in close proximity to the basal surface of the cells of the light organ. The epithelial cells of the light organ are large, the cytoplasm dense, homogeneous and acidophilic. The cytoplasm gives a strong positive reaction for protein. The cytoplasm contains a high density of free ribosomes, patches of dense material, smooth endoplasmic reticulum, glycogen and scattered microtubules. Mitochondria are numerous; they are large, randomly distributed and packed with fine cristae. These cells lack the features characteristic of Malpighian tubule epithelial cells; infolding of the apical and basal cell surfaces is reduced and the cytoplasm contains few organelles. These cells do not contain secretory or photocyte granules and the grainy cell matrix is thought to be the luciferin substrate. Oxygen is supplied via the tracheal layer (which may have secondary reflecting properties) and light production controlled by neurosecretory excitation either directly via synapses, or by hormones. There are no other reports of Malpighian tubules of insects producing light and the fine structure of these cells is distinct. Thus, the swollen distal tips of the Malpighian tubules of the glow-worm undoubtedly constitute a unique luminescent organ.     

Fine structure of the transitional zone of the rat seminiferous tubule

An electron microscopic study was made on the structure of the testicular transitional zone (TZ) in the adult rat. The TZ proper consists of modified Sertoli cellss, with only a few spermatogonia and macrophages, surrounding distally a very narrow lumen. The TZ Sertoli cells have nuclei with a somewhat coarser matrix and more peripheral heterochromatin than Sertoli cell nuclei of the nearby seminiferous tubules, and the electron density of the cytoplasm varies from cell to cell. Smooth endoplasmic reticulum is abundant, but usually there are also scattered ribosomal rosettes and an occasional profile of rough endoplasmic reticulum. Microtubules are very numerous in the columnar portion of the cell, and laminar structures seemingly joining the cell surfaces are sometimes seen. Lipid droplets and lysosmal structures are frequent cellular components in proximal TZ Sertoli cells. Empty intracellular vacuoles are abundant, sometimes arranged around areas of smooth endoplasmic reticulum. Occasionally, membrane-limited fine granules and vacuoles are seen within Sertoli cells and also in the TZ lumen, suggesting a possible secretory activity by these cells. The apical processes of the Sertoli cells form large vacuolar structures, and in the basal parts of the epithelium vacuoles with capillary-like appearance are frequently seen. Phagocytosis of germinal cells by the Sertoli cells occurs in the proximal region of the TZ. Round waste bodies in contact with the Sertoli cell apices protruding into the tubulus rectus, are also common. The tunica propria of the TZ is thickened and somewhat wrinkled, and in the proximal region the myoid cell layer loses its continuity and is replaced by fibroblasts. The epithelium of the tubulus rectus adjacent to the TZ consists of several overlapping epithelial cells. The typical junctional complexes between TZ Sertoli cells appear to be impermeable to the lanthanum tracer.     

Localization of Na+, K+-ATPase to the inside of the basolateral cell membranes of epithelial cells of proximal and distal tubules in rabbit kidney

Summary Cysteine-sensitive alkaline phosphatase and/or ouabain-sensitive Na⁺, K⁺-ATPase were studied by ultrastructure cytochemistry in epithelial cells of proximal and distal kidney tubules. Alkaline phosphatase reactivity was confined to the surface of the microvillous luminal cell membrane of proximal tubule cells, whereas distal tubules and collecting ducts were unreactive. The Na⁺, K⁺-ATPase reactivity was localized evenly along the cytoplasmic side of the basolateral cell membrane of cells of proximal and distal tubules and in collecting ducts. In the proximal tubules, where the activity was strongest, the Na⁺, K⁺-ATPase deposits were also found in the 10–50 nm gap between the cell membrane and the cisternae of tubulo-cisternal endoplasmic reticulum (TER) underlying a major part of the basolateral cell membrane. The restriction of Na⁺, K⁺-ATPase sites, which are involved in extrusion of Na⁺ from the cell, to a narrow cytoplasmic compartment located between the cell membrane and the cisternae of TER, is consistent with a transport role for the TER.     

Specializations of endoplasmic reticulum in bovine placental cells

Summary The endoplasmic reticulum of mononuclear placental cells from 10 cows in different stages of pregnancy has been studied with the electron microscope. Basicaly the cryptal cells are provided with rough surfaced endoplasmic reticulum. In addition, whorls of rough or smooth cytomembranes encircle lipid droplets or plain cytoplasmic matrix. The trophoblastic cells also contain rough surfaced endoplasmic reticulum. Furthermore, skein-like conglomerations of smooth tubules are encountered in some cells. The significance of the membranous structures is discussed.This work was supported by the Swedish Medical Research Council (Project no K 68-12x-2494-01) and NIH General Research Support Grant FR05462.     

Cytological differentiation in the uropygial gland.

Ultrastructural changes were studied in the cells undergoing secretory differentiation in zone I of the tubules of the uropygial gland of White Plymouth Rock chickens. A layer of basal cells and four secretory stages are recognized as the cells migrate from the periphery to the lumen of tubules and progressively elaborate a secretion product. Basal cells, containing rough endoplasmic reticulum and free ribosomes, rest on the basement membrane and are the source from which secretory cells arise. Dilated perinuclear cisternae and the proliferation of smooth endoplasmic reticulum in the form of vesicles, invaginated sacs and cusp-shaped cisternae indicate the onset of lipgenesis in stage I cells. The perinuclear cisternae are more dilated and the endoplasmic reticulum is composed on saccules and cisternae in stage II cells. Stage III cells are characterized by concentric lamellae of endoplasmic reticulum surrounding secretory droplets. Dilated cisternae of endoplasmic reticulum and secretory droplets both contain a reticular substance. The perinuclear cisternae of stage III cells have returned to normal dimensions. Large mature lucent secretory droplets, lined with electron-dense material, fill the cytoplasm ostage IV cells which degenerate and release their secretory product into the tubule lumen. Spherical membrane-bound compartments containing a mottled substance of moderate electron density occur in basal cells and all subsequent secretory stages. These mottled bodies are surrounded by saccules of endoplasmic reticulum in stage II cells and are intimately associated with secretory droplets in stage III cells, but there is no evidence that they give rise to secretory droplets and their role in secretory differentiation is unknown.     

Electron microscopic studies of the epithelial reticular cells of the mouse thymus

Summary Electron microscopic studies have been made of the epithelial reticular cells of the thymus in mice of both sexes ranging in age from 5 to 8 weeks. The epithelial cells generally have long cytoplasmic processes by which they are interconnected and form a network throughout the organ. The processes adhere tightly to one another by desmosomes. At the surface of the organ the processes constitute a thin sheet, and a basement membrane is discernible close and parallel to the free surface of the epithelial sheet. In the cortex the meshes of the epithelial reticulum are filled with numerous lymphoid cells and relatively few mesenchymal reticular cells. The epithelial cells in the cortex are characterized by their slender cytoplasmic processes and by the presence of large round vesicles which contain coarsely granulated, dense material. By the presence of the vesicles as well as desmosomes at junctions of the cytoplasmic processes the epithelial cells can be distinguished from other cells. For comparison the cytological characteristics of the mesenchymal reticular cells are also described. In the medulla two types — reticular and hypertrophic — of epithelial cells are recognized. The cells of reticular type are irregularly stellated in shape with extended cytoplasmic processes. Their cytoplasm often contains considerable amounts of fine filaments in bundles. Due to the relative abundance of free ribonucleoprotein particles and other cytoplasmic components, the cytoplasm appears relatively electronopaque as compared with that of the cells of the other type. The plasma membrane of the cells of reticular type sometimes invaginates into the cytoplasm to enclose a lumen which contains substance of low density and sometimes fine filaments. A basement membrane-like layer is discernible close to the infolded plasma membrane in the lumen. The cells of hypertrophic type are relatively large and round with a few shorter cytoplasmic processes. They are characterized by the abundance of the smooth endoplasmic reticulum which appears as vesicle or sac of small size. These cells often possess peculiar vesicles the wall of which is provided with microvilli projecting into the lumen. Some of these vesicles carry cilia on their wall in addition to the microvilli. The cells of hypertrophic type often undergo degeneration. The degenerating cells are concentrically surrounded by a few neighboring cells of both hypertrophic and reticular types, and Hassall's corpuscles are formed.     

THE FINE STRUCTURE OF TESTICULAR INTERSTITIAL CELLS IN GUINEA PIGS

In guinea pig testes perfused with either glutaraldehyde or osmium tetroxide fixative, the cytoplasm of the interstitial cells contains an exceptionally abundant agranular endoplasmic reticulum. The reticulum in central regions of the cell is a network of interconnected tubules, but in extensive peripheral areas the reticulum is commonly organized into closely packed, flattened cisternae which are fenestrated. Occasional small patches of the granular reticulum occur in the cytoplasm and connect freely with the agranular reticulum. The mitochondria have a dense matrix and contain cristae and some tubules. The Golgi complex is disperse and shows no evidence of secretory material. The cytoplasm also contains lipid droplets. Lipofuscin pigment granules are probably polymorphic residual bodies and contain three components: (1) a dense material which at high magnification shows a 75-A periodicity; (2) a medium-sized lipid droplet; and (3) a cap-like structure. In glutaraldehyde-perfused testis the interstitial cell cytoplasm appears to have the same density from cell to cell, and the agranular reticulum is tubular or cisternal but not in the form of empty vesicles. Thus the "dark" and "light" cells and the vesicular agranular reticulum sometimes encountered in other fixations may be artifacts. Biochemical results from other laboratories, correlated with the present findings, indicate that the membranes of the agranular endoplasmic reticulum in guinea pig interstitial cells are the site of at least two enzymes of androgen biosynthesis, the 17-hydroxylase and the 17-desmolase.     

Observations on the Fine Structure of the Gastric Parietal Cell of the Rat

The structure of the rat parietal cell was examined by electron microscopy. The intercellular and intracellular canalculi are lined by microvilli which are more numerous and larger than those of other gastric cells. The numerous mitochondria have closely packed cristae and a dense matrix containing opaque particles. The cytoplasmic vacuoles typical of parietal cells are part of a network of smooth surfaced tubules and vacuoles (the endoplasmic reticulum) which is intimately associated with the mitochondria and probably connected with the lumen of the canaliculi. Only a few dense particles are found attached to the surface of these tubules. The structure of the parietal cell is compared with that of other cells whose function also is transport of inorganic ions and water. Evidence is presented supporting the hypothesis that parietal cells differentiate from a less structurally specialized cell in the neck region of the gastric gland.     

The secretory apparatus of Ginkgo biloba: structure,differentiation and analysis of the secretory product

Summary The differentiation of the secretory cavities of Ginkgo stem and the structural organization of the epithelial cells were followed by light and electron microscopy. The mode of formation of the cavities is schizo-lysigeneous. Functional complexes of leucoplasts and associated endoplasmic reticulum (ER) membranes are assumed to be the site of synthesis and translocation of the lipophilic secretory product. Most of the endoplasmic reticulum membranes are paired. The content of the cavities was directly collected and analysed by low- and high-resolution mass spectrometry. The cavities contain anacardic acids and cardanols, which are long-chain phenol lipids not characteristic of Ginkgo. The relationship between the plastid/ER complexes and the production of these secondary metabolites is discussed.     

Ultrastructure of the anal organ of Drosophila larva with reference to ion transport

The ultrastructure of the anal organ of the full-grown larva of Drosophila melanogaster is described. The thin cuticle is characterized by epicuticular depressions which contain particulate material. In AgNO(3)-treated larvae, silver grains tend to penetrate the cuticle at the epicuticular depressions. At the basal surface, the epithelial cells exhibit narrow, parallel membrane infoldings which bear a particulate coat on the cytoplasmic surface. The infoldings are also attached around the cytoplasmic surface of endocuticular tubercles, thereby greatly increasing the absorptive surface area. At the apical surface, the membrane invaginations, which are closely associated with mitochondria, anastomose freely and extend deeply into the cytoplasm. The lateral membranes are linked by desmosomes and septate junctions. They are highly folded, are closely associated with mitochondria, and enclose intercellular channels and spaces. The epithelial cells are rich in mitochondria, glycogen particles and tracheoles. Numerous vesicles, multivesicular bodies, lysosome-like dense bodies and sparse endoplasmic reticulum are found in the cytoplasm. In concentrated medium, the epithelial cells show complete absence of the membrane infoldings and invaginations and reduction in the number of mitochondria. The ultrastructural features of the anal organ are consistent with its function in ion transport.     

THE NORMAL FINE STRUCTURE OF OPOSSUM TESTICULAR INTERSTITIAL CELLS

The interstitial tissue of the opossum testis includes interstitial or Leydig cells, macrophages, and small cells which morphologically resemble mesenchymal cells. The latter are thought to give rise to mature interstitial cells. The most prominent feature of the interstitial cell cytoplasm is an exceedingly abundant agranular endoplasmic reticulum. This reticulum is generally in the form of a meshwork of interconnected tubules about 300 to 450 A in diameter, but occasionally it assumes the form of flattened, fenestrated cisternae resembling those of pancreatic acinar cells, except for the lack of ribonucleoprotein particles on the surface of the membranes. The interstitial cells vary considerably in their cytoplasmic density. The majority are quite light, but some appear extremely dense, and in addition usually have a more irregular cell surface, with numerous small pseudopodia. These differences may well reflect variations in physiological state. Cytoplasmic structures previously interpreted as "crystalloids" consist of long bundles of minute parallel tubules, each about 180 A in diameter, which seem to be local differentiations of the endoplasmic reticulum. The mitochondria are rod-shaped, and contain a moderately complex internal membrane structure, and also occasional large inclusions that are spherical and homogeneous. The prominent juxtanuclear Golgi complex contains closely packed flattened sacs and small vesicles. The results of the present study, coupled with biochemical evidence from other laboratories, make it seem highly probable that the agranular endoplasmic reticulum is involved in the synthesis of the steroid hormones produced by the interstitial cell. This finding therefore constitutes one of the first functions of the agranular reticulum for which there is good morphological and biochemical evidence.     

Observations on tubules derived from the endoplasmic reticulum in leaf glands ofPhaseolus vulgaris

Summary Tubular systems present in bean leaf glands have been studied electron microscopically. Ordered arrays of small tubules (290 Å in diameter) arise from the endoplasmic reticulum in early stages of gland development and remain connected to it. Subsequently larger tubules (560–660 Å in diameter) appear among the smaller tubules and gradually replace many of them. The large tubules are not connected to the endoplasmic reticulum. They contain an electron dense material and their walls exhibit a patterned substructure. In older gland cells the bundles of large tubules run randomly through the cytoplasm. The relationship of the two types of gland tubules to conventional microtubules has been examined morphologically and experimentally. The small tubules have larger diameters and thicker walls than microtubules. Neither type of gland tubule is affected by low temperature or colchicine, or, in thin sections, by pepsin digestion. This suggests that these tubules are not closely related chemically to either cytoplasmic or ciliary microtubules. The two systems of tubules are closely associated with prominent protein vacuoles in the gland cells, but are not directly connected to them.This work was supported in part by grant no. GB-6161 from the National Science Foundation.     

Structure and development of the chloroplast in Chlamydomonas. I. The normal green cell

The cytoplasmic organization of a normal green strain of the alga Chlamydomonas reinhardi has been investigated with the electron microscope using thin sections of OsO(4) fixed material. The detailed organization of the chloroplast has been of special interest. The chloroplast, a cup-shaped organelle, surrounded by a double membrane, consists of: (1) discs about 1 micron in diameter, considered to represent the basic structural unit of the chloroplast, and each composed of a pair of membranes joined at their ends to form a flat closed vesicle; the discs are grouped into stacks resembling the grana of higher plants; (2) matrix material of low density in which the discs are embedded; (3) starch grains; (4) the pyrenoid, a non-lamellar region associated with starch synthesis, and containing tubules which connect with the lamellae; (5) the eyespot, a differentiated region containing two or three plates of hexagonally packed, carotenoid-containing granules, located between discs, and associated with phototaxis. In addition to the chloroplast, the cytoplasm contains various membranous and granular components, including mitochondria, endoplasmic reticulum, and dictyosomes, identified on the basis of morphological comparability with structures seen in animal cells. The nucleus, not investigated in detail in this study, contains a large, granular nucleolus and is surrounded by a nuclear envelope which is provided with pores and exhibits instances of continuity with the endoplasmic reticulum of the cytoplasm.     

Aspect ultrastructural de la lipogénèse dans le tissu adipeux brun

Summary In interscapular brown fat of the rat, appropriately processed so as to maintain membrane structures intact, lipid droplets are in fact liposomes, i.e., lipid filled vacuoles surrounded by a membrane that is related to the smooth endoplasmic reticulum. Thus the endoplasmic reticulum appears as an important component of brown fat cells. — After complete lipid depletion has been achieved by 7 days of fasting, feeding with glucose results in sudden and very conspicuous increase of smooth endoplasmic reticulum; dilatations of the perinuclear cistern and pinocytotic activity at the periphery of the cell contribute to this increase. Simultaneously the cytoplasmic matrix is heavily loaded with glycogen in particulate form. Lipogenesis, as far as can be appreciated by different degrees in electron density, takes place inside the vesicles of the endoplasmic reticulum; increase of such small lipid vacuoles then leads to reconstitution of liposomes; return to the normal aspect is completed 12 hours after the beginning of feeding. — During the phase of glycogen overloading and resulting lipogenesis, glycogen particles may be found in intercellular and pericapillary spaces; the significance of this finding is discussed. — The fundamental difference between both types of fat cells seems to be concerned with the site of lipogenesis: this takes place in the cytoplasmic matrix of white fat cells, so that lipid droplets aggregate without any limiting membrane, whereas in brown fat cells lipogenesis occurs inside cavities of the endoplasmic reticulum and lipids remain permanently enclosed in membranes. This process appears similar to what may be observed occasionally in liver and normally in adrenal cortex, and this might presumably lead to a physico-chemical understanding of the particular aspect of lipogenesis in brown fat, as compared to that in common white fat.

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Travail dédié au Professeur W. Bargmann, en témoignage d'admiration, à l'occasion de son 60e anniversaire.     

Microlipid droplets in milk secreting mammary epithelial cells: evidence that they originate from endoplasmic reticulum and are precursors of milk lipid globules

Microlipid droplets, structures with diameters less than 0.5 micron, resemble larger cytoplasmic lipid droplets of milk secreting mammary epithelial cells in triacylglycerol core and surface coat composition. Previously, evidence was obtained that microlipid droplets fuse with and support growth of cytoplasmic lipid droplets, which are immediate precursors of large milk lipid globules. Morphological observations suggested that microlipid droplets may also be secreted directly from mammary epithelial cells, yielding the very small lipid globules of milk. The secretion mechanism, which involves envelopment of triacylglycerol droplets in apical plasma membrane, appeared to be the same for microlipid droplets as for larger cytoplasmic lipid droplets. Microlipid droplets appeared to originate by blebbing from cisternae of endoplasmic reticulum. By immunogold cytochemical localization and by immunological identification of electrophoretically separated polypeptides, endoplasmic reticulum, micro- and cytoplasmic lipid droplets, and milk lipid globules had a number of common polypeptides. Kinetics of incorporation of radiolabeled palmitate or glycerol into triacylglycerols and phospholipids were consistent with a possible endoplasmic reticulum origin of microlipid droplets and with the view that microlipid droplets may be secreted directly from the cell or may fuse with cytoplasmic lipid droplets.     

THE EFFECTS OF ENUCLEATION ON THE CYTOPLASMIC MEMBRANES OF AMOEBA PROTEUS

The dependence of cytoplasmic membranes upon the nucleus was studied by examining enucleated amebae with the electron microscope at intervals up to 1 wk after enucleation. Amebae were cut into two approximately equal parts, and the fine structure of the enucleated portions was compared with that of the nucleated parts and starved whole cells which had been maintained under the same conditions. Golgi bodies were diminished in size 1 day after enucleation and were not detected in cells enucleated for more than 2 days. The endoplasmic reticulum of enucleated cells appeared to increase in amount and underwent changes in its morphology. The sparsely scattered short tubules of granular endoplasmic reticulum present in unmanipulated amebae from stock cultures were replaced in 1–3-day enucleates by long narrow cisternae. In 3–7-day enucleates, similar cisternae of granular endoplasmic reticulum encircled areas of cytoplasm partially or completely. It was estimated that in most cases hundreds of these areas encircled by two rough membranes were formed per enucleated cell. The number of ribosomes studding the surface of the endoplasmic reticulum decreased progressively with time after enucleation. In contrast, the membranes of nucleated parts and starved whole cells did not undergo these changes. The possible identification of membrane-encircled areas as cytolysomes and their mode of formation are considered. Implications of the observations regarding nuclear regulation of the form of the Golgi apparatus and the endoplasmic reticulum are discussed.