生殖生理学Chapter4

CHAPTER 4 HISTOLOGICAL PHYSIOLOGY OF REPRODUCTION FEMALE REPRODUCTIVE ORGANS

Ovary

The ovary consists of an outer zone, the cortex, and an internal zone, the medulla (Baker, 1972). The germinal epithelium on the ovarian surface is cuboidal (Fig.4-1). The ovarian follicles are in different stages of development (Table 4-1).

Follicular Cells. In order to remain alive and viable, the germ cells must be enclosed by fol-licle cells. Contact between the oocyte and the follicle cells is maintained within the developing ovary, even though the deposition of the zona pellucida induces a topographic separation (Fig.

4-2). The follicle cell and the oocyte are connected by cytoplasmic processes that originate from the follicle cell and contain mitochondria, lipid droplets and pleomorphic bodies reminiscent of lysosomes. These projections terminate in either the perivitelline space or at the oolemma (Fig.

4-3). While the projections within the perivitelline space degenerate, the other projections become surrounded by oocyte microvilli and are connected to the oocyte.by desmosomes.

Although the association of the oocyte with follicle cells is essential for the maintenance of the oocyte, follicle cell contact does not ensure the survival of the oocyte because oocytes within cystic follicles are enclosed by follicle cells but are atretic. In addition, grossly degenerative oocytes are found within apparently "healthy" antral follicles, which suggests that the oocyte deteriorates prior to the follicle.

The functions of granulosa cells and the maturational process of the oocyte are coordinated by intercommunication between cells within the membrana granulosa and between granulosa cells and the oocyte. The integrity of the junctional complexes may be estrogen-dependent, thus indicating a possible intrafollicular relationship between the steroidogenic function of the thecal cells and the maturation of the oocyte.

Degeneration of Oogonia. Oogonia have three phases of degeneration: phase 1--chromatin condenses at the periphery of the nucleus with no effect on the cytoplasmic organelles; phase 2--a dense ring of chromatin develops at the margin of the nucleus and the cytoplasmic organelles swell and rupture, resulting in cytoplasmic vacuolization; phase 3--granulosa cells often invade and subsequently phagocytie the atretic oogonia.

Corpus Luteum. After ovulation, the corpus luteum differentiates fully into a typical endo-crine gland, secreting progesterone. The steroid hormone secreted by the corpus luteum has sev-eral functions: (a) it prepares the endometrium for implantation: (b) it enables the blastocyst to implant, and (c) it maintains pregnancy after implantation (Hansel et al., 1973). The functional lifespan of the corpus luteum, which varies with the species, depends primarily upon the fate of the embryo. Following implantation of the blastocyst, the corpus luteum undergoes extensive enlargement and continues to synthesize and secrete steroid hormones (the corpus luteum of pregnancy).

Female Reproductive Tract

Remarkable differences exist in the pattern of tissue organization of surface epithelium of different segments of the reproductive organs (Hafez, 1978a,b). In most organs, the cells are uniform in size, closely packed, and resemble cobblestones（鹅卵石）. In some instances, the surfaces of the cells are ill-defined and covered with short microvilli and residual mucus.

Table 4-1 Functional histology of the mammalian ovary

Anatomic Functional Unit Histologic Characteristics

Superficial epithelium Tunica albuginea

Ovarian cortex

Ovarian medulla Ovarian stroma Smooth muscle

Ovarian Follicles Primary follicle

Growing follicle Secondary follicle

Tertiary (vesicular) follicle

Graafian follicle Preovulatory follicle A surface layer of flattened epithelium (commonly and incorrectly known as germinal epithelium) with abundant microvilli.

Dense fibrous connective tissue covering the whole ovary just beneath the superficial epithelium. Connective tissue cells near the surface are arranged roughly parallel to the ovarian surface and are somewhat denser than the cells lying toward the medulla.

Contains several primary follicles (with oocytes in a quiescent state) and a few large follicles. During each estrus cycle, a variable number of follicles undergo rapid growth and development, culminating in the process of ovulation.

Loose connective tissue that contains nerves, lymphatics, and tortuous thin-walled blood vessels, collagen and elastic fibers, fibroblasts and scattered smooth muscle cells.

Poorly differentiated, embryonal-mesenchymal-like cells capable of undergoing complex morphologic alterations during the reproductive life; stromal cells can give rise to theca interna cells or interstitial cells.

Smooth muscle cells are present throughout the ovary, especially in the cortical stroma where they intermingle with the cells of the theca. Ovarian myoid cells are similar to smooth muscle cells of other tissues. These cells contain large numbers of microfilaments arranged in characteristic bundles and also micropinocytotic vesicles located just beneath the plasmalemma. The cholinergic receptors on smooth muscle cells may mediate the contraction of the graafian follicle. Thus, smooth muscle cells and neural elements may be directly involved in ovulation. The presence of smooth muscle cells, especially in the perifollicular regions, may be involved in "squeezing the follicle" during ovulation.

An oocyte enclosed by a single layer of flattened or cuboidal follicular cells and surrounded by interstitial tissue.

Oocyte with increased diameter and increased number of layers of follicular cells; zona pellucida is present around the oocyte. Flattened granulosa cells of the primordial or unilaminar follicle proliferate, assuming an irregular, polyhedral appearance.

Under the influence of pituitary gonadotrophin, the granulosa cells of multilayered follicles secrete a fluid, the liquor folliculi, which accumulates in the intercellular spaces. The continued secretion and accumulation of liquor folliculi result in the dissociation of granulosa cells, which causes the formation of a large, fluid-filled cavity-the antrum. The zona pellucida is surrounded by a solid mass of radiating follicular cells, forming the corona radiata. Follicular cells increase in size; the antrum is filled with follicular fluid (liquor folliculi). The oocvte is pressed to one side, surrounded by an accumulation of follicular cells (cumulus oophorus); elsewhere in the follicular cavity an epithelium of fairly uniform thickness called the membrana granulosa has formed. The theca interna and theca externa have formed.

A blister-like structure protruding from the ovarian surface due to rapid accumulation of follicular fluid and thinning of the granulosa layer. The viscous liquor folliculi is formed from the secretions of granulosa cells and plasma proteins transported into the follicle by transudation. The cumulus oophorus is detached from the thinned and extensivelv dissociated stratum granulosum.

Corpus hemorrhagicum Corpus luteum Corpus albicans The oocyte lies free in the liquor folliculi, surrounded by an irregular mass of cells. Dramatic changes are noticeable at the subcellular level, particularly in the Golgi complex, which is involved in the formation of the zona pellucida and cortical granules. The oocyte, in the prophase of meiosis (dictyate stage), resumes meiosis several hours before ovulation. The first meiotic (maturational) division is associated with extrusion of the first polar body, which may briefly remain attached to the oocyte by a cytoplasmic bridge.

The follicular cavity is filled with lymph and blood from broken thecal vessels, blood from follicular fluids and blood from small vessels that hemorrhaged at ovulation. It acts as a "stopper," sealing the residual cavity after discharge of oocyte. Intact vessels and connective tissue cells from the surrounding theca begin to proliferate.

The transformation of a ruptured follicle into a corpus luteum involves characteristic folding of the granulosa layer toward the central portion of the residual cavity; luteinization of the granulosa cells occurs under the influence of LH. Lutein cells, polyhedral in form or without definite cell walls, are arranged in irregular masses. The cytoplasm may be clear or granular according to the secretory and functional activity.

White fibrous tissue that forms from the corpus luteum of previous ovulations.

The oviduct, uterus and cervix are composed basically of secretory cells and ciliated cells. The secretory cells secrete the luminal fluids necessary for the transport and nutrition of sper-matozoa and eggs and for cleavage of fertilized eggs and development of preimplantation blas-tocysts.

Ciliated cells are found singly or in groups arranged in rows or a mosaic pattern. Cilia first grow at the periphery of the cell and then fill in the central area. Cilia appear first as stubby (short and thick), short cylinders on those cells that have large numbers of microvilli on their surfaces.

In the female reproductive tract, kinocilia play an important role in the transport of particles and in directing the flow of luminal fluids. Two types of ciliary motility have been recognized: an effective stroke and a recovery stroke. In the effective stroke the cilia sweep rapidly in a still and slightly wavy motion. In the recovery stroke the cilia bend near the basal body and the degree of bending proceeds as a slow wave toward the tip. The recovery stroke carries the cilia back to the effective stroke position. Cilia beat approximately 1200 times/minute. The kinocilia in the female reproductive tract beat rhythmically toward the vagina creating a directional flow of luminal fluids.

Oviduct

The wall of the oviduct is composed of mucosa, muscularis and serosa (Hafez and Blandau, 1969).

Oviductal Mucosa. The oviductal mucosa is made of primary, secondarv and tertiary folds (Figs.4-4 and 4-5). The mucosa in the ampulla is thrown into high, branched folds that decrease in height toward the isthmus and become low ridges in the uterotubal junction. The complex arrangement of these mucosal folds in the ampulla almost completely fills the lumen so that there is only a potential space. Fluid is at a minimum; thus, the cumulus mass is in the intimate contact with the ciliated mucosa.

The mucosa consists of one layer of columnar epithelial cells. The underlying suhmucosa of smooth muscle fibers and connective tissue is permeated by fine blood and lymph vessels. The

epithelium contains ciliated and nonciliated cells, together with "peg cells" (Stiftzellen or intercalary cells), which are presumably depleted secretory cells ( Fig. -1-6). The peg cells are long, rod-like slender cells with a dark compressed nucleus and luminal surface indentations with practically no cytoplasm. They are usually located in the basal portions of the folds near secretory cells and are less common at the uterine end of the oviduct. The oviductal epithelium undergoes morphologic and cytologic changes during the estrous cycle.

Ciliated Cells. The ciliated cells of the oviductal mucosa have a slender motile cilia (kinocilia) that extend into the lumen (Fig. 4-6). The cilia conform to the standard structural plan of kinocilia, with nine pairs of fibrils arranged concentrically around another central pair. Ciliated cells are characterized by a fine granular cytoplasm containing vesiculated endoplasmic reticulum, large mitochondria and cytoplasmic droplets. Large, centrally placed, round or oval nuclei are surrounded by a perinuclear halo（晕轮）of clear cytoplasm.

The apical surfaces of both ciliated and secretory cells are covered by microvilli that are uni-formly distributed. Microvilli found on secretory cells tend to be long and slender; on ciliated cells, these surface modifications are usually hidden by cilia in many histologic sections because of the angle of the section.

The rate of beat of cilia is affected by the levels of ovarian hormones, activity being maximal at ovulation or shortly afterward when the stroke of the cilia in the fimbriated portion of the ovi-ducts is closely synchronized and direction-oriented toward the ostium. The action of ciliary beat seems to enable the egg within the surrounding cumulus cells to be stripped from the surface of the collapsing follicles toward the ostium of the oviduct. The percentage of ciliated cells decreases gradually in the ampulla toward the isthmus, and reaches a maximum in the fimbriae and infundibulum. Ciliated cells are noted in large numbers at the apices of the mucosal folds. Variations in the percentage of ciliated and secretory cells along the length of the oviduct have some functional significance. Ciliated cells are most abundant where the egg is picked up from the ovarian surface, whereas secretory cells are abundant where luminal fluids are needed as a medium for the interaction of eggs and spermatozoa.

Although the ciliated cells in the ampulla appear to arise from crypt-like crevices because of the apical protrusion of the nonciliated cells, the ciliary tips have free access of movement into the lumen. In the isthmus there are a few ciliated cells located deep between the tall secretory ones. The tips of their cilia hardly reach the oviductal lumen and thus exert no effect on egg transport. The cilia beat toward the uterus. Their activity, coupled with oviductal contractions, keeps oviductal eggs in constant rotation, which is essential for bringing egg and sperm together (fertilization) and preventing oviductal implantation.

Ciliation of the oviduct is hormonally controlled in the rhesus monkey: cilia disappear almost completely after hypophysectomy and develop in response to administration of exogenous estrogens. The oviducts atrophy and deciliate during anestrus, hypertrophy and become reciliated during proestrus and estrus, and atrophy and deciliate during pregnancy.

Infections of the female reproductive tract are associated with dramatic changes in cell mor-phology. Infection is usually associated with the loss of ciliated cells. Inflamed oviductal epithe-lium has ciliated cells devoid of cilia that retain a certain number of ciliary basal bodies, the pre-cursors of ciliary shafts. A decrease in the number of cilia may lead to the accumulation of tubal fluid and inflammatory exudate, which contributes to the agglutination of tubal plicae and sub-sequent development of salpingitis.

Secretoy Cells and Fluid. The secretory cells of the oviductal mucosa are nonciliated and characteristically contain secretory granules, the size and number of which vary widely among species and during different phases of the estrous cycle. The apical surface of the nonciliated cells is covered with numerous microvilli (Figs.4-6, 4-7). The structure of the secretory cells, the epithelial height and the secretory activity change throughout the estrous cycle, with maximal changes occurring near the time of ovulation. Secretory granules accumulated in epithelial cells during the follicular phase of the cycle are released into the lumen after ovulation, causing a reduction in epithelial height. These variations generally become less pronounced moving from the infundibulum to the isthmus. There are quantitative and qualitative variations in the secretory granules in the same cell during different stages of the reproductive cycle. The release of secretory granules from the nonciliated cells of the oviducts is programmed by gonadal hormones.

Definite cytologic changes during the follicular phase precede secretory activity. The en-doplasmic reticulum of the secretory cells spreads out, the mitochondria swell as their matrices fill with a granular substance, and the Golgi apparatus becomes well developed. During the luteal phase, numerous secretory droplets appear, the endoplasmic reticulum dilates, mitochondria become fewer and the Golgi apparatus expands.

The metabolic activity of the epithelium of the oviduct appears to be most pronounced in the vicinity of the egg. In the rabbit, for example, the uptake of radioactive sulfate is always highest in the segment that contains the egg as it passes along toward the uterus.

By means of extra- and intra-abdominal devices that are used to cannulate the oviduct and collect its fluids, it has been possible to show that ovarian hormones also regulate the secretory activity of the epithelium of the oviduct (Perkins et al., 1965). The secretions are composed predominantly of mucoproteins and mucopolysaccharides.

Muscularis and Serosa. The tunica muscularis consists of an inner circular layer and an outer longitudinal layer of smooth (unstriated) muscle (Fig.4-8). The contractility of oviductal musculature, under hormonal control, regulates in part the transport of eggs and spermatozoa.

Uterus

The uterus consists of a thin outer layer, the perimetrium, a thick myometrium composed of inner circular and outer longitudinal smooth muscle layers, and an inner layer, the endometrium. The changes that occur in the endometrium during the estrous cycle prepare the uterus to receive the blastocyst and therefore play a major role in the reproductive process.

Endometrium. The endometrium is a highly glandular structure consisting of an epithelial lining of the lumen, a glandular layer and connective tissue. It varies in thickness and vascularity with changes in ovarian hormones throughout the estrous cycle and pregnancy.

The columnar epithelial cells that line the uterine lumen play an important role in the in-teraction between the blastocyst and endometrium during the initial stages of pregnancy. To facilitate the attachment of the blastocyst, the Luminal epithelial cells undergo structural changes that are regulated by progestins (progesterone) and estrogens.

The endometrium is characterized by the presence of numerous openings of the endometrial glands (Figs. 4-8, 4-9). Ciliated cells are less abundant in the endometrium than the oviductal epithelium. Large cytoplasmic projections are noted at the apical membrane of the nonciliated cells. The abundance, length, shape and interbranching of apical microvilli vary throughout the

cycle. The development of apical microvilli, the synthesis, storage and release of endometrial secretory granules and ciliogenesis are hormone-dependent. Degenerated cells are found at random at different stages of the estrous cycle. Following the release of secretory material from the protruding cell membrane, the cell collapses and becomes wrinkled and devoid of microvilli.

Caruncles. The inner surface of the uterus in ruminants contains nonglandular projections, the caruncles. They are arranged in four rows, extending from the uterine body into the two uterine horns, and consist of connective tissue comparable to that found in the cortical stroma of the ovary. The deeper areas between the prominences are rich in blood vessels but contain no glands.

The uterus of the nonpregnant cow has 70 to 120 caruncles, each measuring approximately

15 mm in diameter. During pregnancy, they may attain a diameter of 10 cm and appear spongy due to the numerous crypts that receive the placental chorionic villi. These villi develop in lo-calized areas, the cotyledons, which invade the caruncles. The cotyledons and caruncles are re-ferred to together as the placentome.

In horses and swine, the uterus has no caruncles. Instead, the mucosa is characterized by conspicuous longitudinal folds that pass into the cervix to form the internal and external orifices.

Endometrial Glands. The endometrial glands are simple branched tubular glands that are more or less coiled, especially toward their ends. The glands possess a lamellar connective tissue and are lined by simple or ciliated columnar epithelium.

These glands are scattered throughout the endometrium with the exception of the caruncles. They are branched, tubular and coiled, especially toward the ends. Their density varies with the species, breed, parity and estrous cycle. The variation in proximity to one another during the estrous cycle is largely a result of changes in diameter and amount of stromal ground substance. The number of glands is higher in the horns than in the mucosa adjacent to the cervix. The glands may be rapidly increased by budding and outward growth from the basal zone. Buds that do not reach the surface become dilated and cystic.

The cyclic shedding of the endometrial surface epithelium followed by regeneration causes no extensive bleeding, such as that observed in primates in which a substantial portion of the en-dometrium is shed. During "metestrus bleeding" in the cow, the caruncles show a pronounced capillary distention, but the epithelium remains intact.

Myometrium. The myometrium is the muscular portion of the uterine wall. It consists of two layers of smooth muscle: a thick inner circular layer and a thinner outer longitudinal layer. Between them lies a vascular layer made of blood and lymph vessels, nerves and connective tissue. This layer is not distinct in the sow and mare, but may even occur within the circular layer in ruminants. During pregnancy the amount of muscle tissue in the uterine wall increases, both by cell enlargement and cell number.

The serosa (perimetrium), the stratum musculare and the layer of longitudinal muscle are all continuations of the broad ligament, which invests and suspends the uterus.

Cervix

The cervix is a sphincter-like structure that projects caudally into the vagina. It is charac-terized by a thick wall, a constricted lumen, and transverse or spirally interlocking ridges that de-velop to varying degrees in the different species. These ridges are especially prominent in rumi-

nants, in which they fit into each other to close the cervix securely.

Cervical Mucosa. The cervix does not contain any glands, as is assumed erroneously. The cervical mucosa is thrown into primary and secondary mucosal crypts that provide an extensive secretory surface. These intricacies give the cervix its typical "fern leaf" microscopic appearance. The cervical mucosa is made of two types of columnar epithelial cells: ciliated cells with kinocilia and nonciliated secretory cells (Fig. 4-10). The kinocilia beat toward the vagina and the nonciliated cells contain massive numbers of secretory granules. The greatest secretory activity of these cells occurs at estrus (Fig. 4-11).

The cervical crypts are more developed in the cow than in other farm animals. The mucosa lies in large folds, which are tallest and thickest in the cow and thinnest in the ewe. The folds vary in height, particularly within the cow and goat cervix. In all three species, the cervical folds are taller, more numerous and less regular in size in the cranial part of the cervix than in the caudal part (Fig. 4-12).

At estrus, the cells in the bases of the folds and the secondary indentations act as if they are more responsive to estrogenic stimulation than the cells near the lumen (Heydon and Adams, 1979). This has implications for sperm transport because the mucus coming out of cells is believed to guide spermatozoa along the "lines of strain" down to the mucosa near its origin. A greater mucous response to estrogen in the bases of the folds and secondary indentations will result in more sperm being guided specifically into these areas, thus aiding in the physical entrapment of spermatozoa. There is evidence that the biophysical characteristics of cervical mucus vary in various parts of the cervical crypts. Using histochemical techniques, sulfomucins occur in cells of the cervical folds closest to the lumen, while in the cells in the crypts or the secondary indentations mucus is more abundant and stains for sialomucins (Heydon and Adams, 1979). The epithelium of the cervix of ovariectomized females contains only a small amount of mucus, which stains for sulfomucin. Treatment with estrogen produces large amounts of mucus in the cells, and this stains a sialomucin.

Cervical Wall. The cervical wall consists primarily of fibrous, elastic and collagenous tissue and a small amount of muscle. The connective tissue is made of fibrous constituents, cellular components and ground substance. Collagen, the principal fibrous element, has a high tensile strength. Collagenase, a speciflc degenerative enzyme for collagen, may be involved in the process of labor. The reticular network is an intimate part of the collagen system. The mass of the cervix increases as gestation advances and, although involution occurs after parturition, part of the gain in mass is maintained.

Vagina

The normal squamous epithelium of the lower vagina is smooth, with little undulation of the surface architecture. The cells are flat, polygonal, with demonstrable but thin-edged interdigitating borders. The cells, in multiple layers, overlay each other irregularly, much like haphazardly (randomly) layered shingles（鹅卵石）on a roof. The edges of these cells roll back, giving the characteristic picture of exfoliation as they lift up from their borders. The exfoliated cells appear wrinkled as if dried up; with lost or obscured surface microridges.

The surface of the vaginal cell is made of numerous microridges that run longitudinally or in circles ( Fig. 4-13). In this multilayered stratified epithelium, the cells are wedged one upon the

other by interlocking opposed microridges, thus forming a firm surface. The morphology and pat-tern of these microridges, which affect the firmness of the epithelium, vary throughout the re-productive cycle. The microridges exhibit a regular pattern during pregnancy, while pores appear within the microridges of the cells during the estrous cycle.

Sequential changes in the cell types present in the vaginal smears are unreliable as the sole means of detecting ovulation in ewes.

Table 4-2 Functional histology of the testis

Segment

Tunica albuginea Seminiferous tubules

Spermatogonium Primary spermatocytes Secondary spermatocytes Spermatids

Sertoli cells A thick, white capsule of connective tissue surrounding the testis; made primarily of interlacing series of collagenous fibers with elongated and flattened nuclei of the fibroblasts.

Appear as large isolated structures, round or oblong in outline; varying appearance due to the complex coiling of the tubules at many different angles and levels. Between the tubules are blood vessels of variable diameter (smaller than tubules) packed with erythrocytes and embedded in small masses of interstitial (Ieydig) cells, which produce the male sex hormones. Strands of connective tissue form a thin layer around each tubule.

Lie in the outermost region of the tubule; round nuclei appear as an irregular layer just within surrounding connective tissue. Nuclei may be recognized by small size and dark stain due to presence of large numbers of chromatin granules. Located just inside an irregular layer of spermatogonia and Sertoli cells; nuclei are noticeably larger than those of the spermatogonia and stain lighter, although they still contain a considerable amount of granular chromatin.

Maturation divisions and secondary spermatocytes are not seen in the average tubule owing to the short duration of these stages.

Located internally to primary spermatocytes. Cells are small and round with light-staining nuclei. Layer of spermatids may be several cells in thickness. Spermatozoa lie along the border of the lumen; their long tails extending into the cavity as filamentous structures, their heads, appearing as dark-staining small points or short lines. The sperm heads are lodged in deep indentations of the surface of the Sertoli cell, but never actually within the cytoplasm.

Large and relatively clear except for the prominent, dark-staining nucleolus. Cytoplasm is diffuse and its limits are indefinite.

Vulva

The vulva, made of the labia majora and minora, the clitoris and the vaginal introitus (inlet), is covered by stratified squamous epithelium similar to all other skin areas, with some follicles and sebaceous and sweat glands. The surfaces of the cells show the usual interlacing microridges. The edges of these cells are thin in contrast to the slightly raised terminal bars on the squamous cells of the upper vagina.

MALE REPRODUCTIVE ORGANS

Extensive investigations have been conducted on the anatomy, physiology and biochemistry of the mammalian testis (Johnson et al., 1970), epididymis (Martan, 1969) and male accessory or-gans (Spring-Mills and Hafez, 1979, 1980).

The histologic and cytologic characteristics of the cellular components of the seminiferous tubules are summarized in Table 4-2 and illustrated in Figures 4-14 and 4-15. While in the epididymis, the spermatozoa, helped by the secretions of the epithelium, approach full maturity

and increase their motility. They move through the duct from the caput epididymidis to the cauda epididymidis where they are transported to the vas deferens.

The histologic characteristics of the male accessory organs are illustrated in Figures 4-14 and 4-15. Secretions are stored in the lumen of the male accessory glands. Following the emptying of the glands, there is a continuous accumulation of secretion in the lumen. The accessory glands are continuously active in synthesizing and secreting protein, the secretion either being used during copulation or lost by spontaneous ejaculation.

Epididymis

The eff'erent ducts connect the rete testis to the epididymal duct, which follows a continuous but highly convoluted course. The epididymis, more than 120 feet long in the bull and about 180 feet long in the boar, is divided macroscopically into three gross regions: head (caput or initial), body (corpus or middle) and tail (cauda or terminal). There is a progressive decrease in the height of the epithelium and stereocilia and a widening of the lumen throughout the three segments. The first 2 segments are concerned with sperm maturation whereas the terminal segment is for sperm storage. The epithelial lining of the epididymis consists of different kinds of cells: principal columnar cells, small basal cells on basal membrane and some lymphocytes.The lumen of the epididymal tubules is lined with epithelium made of a basal layer of small cells and a surface layer of tall columnar ciliated cells. Masses of sperm are often found in the lumen. Spaces between tubules are filled with a loose connective tissue.

Vas Deferens

The mucosa is thrown into longitudinal folds. Near the epididymal end of the vas the epithe-lium resembles that of the epididymis: the nonciliated cells have little secretory activity. The lumen is lined with pseudostratified epithelium that may or may not be stereociliated. The height of the epithelium is lower than that in the epididymis. The musculature consists of an inner and outer longitudinal layer and a thicker middle circular layer. The three layers are not easily recognizable because muscle fibers of the layers are interwoven. There are species differences in the thickness of the various muscle layers, the length of the duct and the presence or absence of the ampulla.

Male Accessory Organs

Ampullary Gland (Ampulla Ductus Deferentis). These glands open into the urethra. The vas deferens forms a thickening called the ampulla; this thickening is due to an increase in the size of the mucosa, which possesses branched tubular glands with sac-like dilations. The gland consists of tubules enclosed by a capsule of connective tissue and smooth muscle. Strands of connective tissue extend from the capsule to form septa between the tubules. Smooth muscle cells appear in the anterior septa but disappear as the duct approaches the urethra. The ampullae, absent in the boar, are highly developed in the stallion and contribute ergothioneine（麦角硫因）—a

sulfur-containing nitrogenous base—to the ejaculate.

Seminal Vesicles (Vesicula Seminalis). The seminal vesicle originates as an evagination of

the vas deferens. Each consists of an elongated simple or branched tubule that is distended with fluid during the breeding season. The gland is composed of an inner secretory epithelium, a middle layer of smooth muscle, and an external layer of collagenous and elastic connective tissue. The epithelium is pseudostratifled or columnar. Folds of mucous membrane unite to form a reticulated surface. Vesicles may contain a granular secretion that stains strongly with eosin (red). The muscular walls consist chiefly of circular muscles. The seminal vesicles of the bull have numerous lobes. In the stallion, elongated pear-shaped glands contribute the gel to the ejaculate. In the boar, the well-developed seminal vesicles are filled with a milky, highly viscous fluid. The dog and cat have no seminal vesicles.

In studies on the blood-brain barrier performed in the early 1900s, the concept of a

blood-testis barrier arose with the demonstration that certain dyes introduced into the bloodstream were excluded from the seminiferous tubules of the testis. Subsequently, physiologic studies demonstrated that proteins, amino acids and certain ions in plasma were also largely excluded from the seminiferous tubules (Hoffer and Hinton, 1984).

Prostate Glands (Glandula Prostata). The prostate gland is present in two forms: (1) as a mass of tissue located at the vas deferens-urethral junction; this type is encapsulated and lobulated to varying degrees; it penetrates the urethral muscularis and opens into the urethra by means of one pair or more ducts: (2) as a disseminate type, which extends along the entire prostatic and most of the membranous urethra.

In the ram and the goat, the prostate is a primitive disseminate type and consists of glands that do not penetrate muscle surrounding the pelvic part of the urethra. In the bull and boar, the prostate resembles both the lobed and disseminate type. In general, the prostate is a tubulo- alveolar gland with epithelia folds extending into the lumen. Spherical secretory granules are released from the free surface, particularly during periods of high sexual activity.

Bulbourethral Glands (Cowper's Glands, Glandula Bulbourethralis). These glands are round, compact bodies except in the boar, in which they are large, almost cylindrical and filled with a viscous, rubber-like, white secretion that forms a gel in the ejaculate. They are located dorsally to the bulb of the penis, partially or completely embedded in the bulbocavernous muscle. The compound tubulo-alveolar glands resemble mucous glands and are surrounded by a capsule of connective tissue and striated muscle. Connective tissue forms septa and trabeculae that subdivide the glands.

《组织学与胚胎学》同步练习题——女性生殖系统(含答案,人卫版)

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第十二章生殖 一、名词解释 1、生殖 2、月经 3、排卵 4、妊娠 5、分娩 二、填空题 三、选择题 1、精子的发育过程正确的是（ A ） A、精原细胞→初级精母细胞→次级精母细胞→精子细胞→精子 B、初级精母细胞→精原细胞→次级精母细胞→精子细胞→精子 C、精原细胞→初级精母细胞→精子细胞→次级精母细胞→精子

D、精原细胞→精子细胞→初级精母细胞→次级精母细胞→精子 E、精原细胞→次级精母细胞→精子细胞→初级精母细胞→精子 2、体内精子储存在 D A、睾丸 B、前列腺 C、精囊腺 D、附睾和输精管 E、尿道球腺 3、睾酮的主要产生部位是 B A、睾丸生精细胞 B、睾丸间质细胞 C、睾丸支持细胞 D、曲细精管上皮细胞 E、肾上腺皮质网状带细胞 4、下列哪项不属于睾酮的生理作用 C A、维持生精作用 B、刺激生殖器官生长发育 C、促进乳腺发育 D、促进蛋白合成 E、促进男性副性征出现 5、排卵后形成的黄体可分泌 D A、孕酮 B、黄体生成素 C、卵泡刺激素 D、雌激素和孕酮 E、黄体生成素和孕酮 6、妊娠时维持黄体功能的主要激素是 E

A、雌激素 B、孕激素 C、卵泡刺激素 D、黄体生成素 E、绒毛膜促性腺激素 7、血液中哪种激素出现高峰可以作为排卵的标志 C A、雌激素 B、孕激素 C、黄体生成素 D、卵泡刺激素 E、卵泡刺激素释放激素 8、下列哪项不属于雌激素的生理作用 B A、使卵泡发育成熟、排卵 B、使子宫内膜发生分泌期变化 C、使输卵管运动增强 D、刺激阴道上皮细胞增生、角化 E、促进乳腺发育 9、关于孕酮的生理作用的叙述,下列哪项是错误的 E A、使子宫内膜呈分泌期变化 B、使子宫肌活动减弱 C、抑制母体免疫排斥反应 D、促进乳腺腺泡发育 E、使排卵后基础体温降低 10、女性基础体温在排卵后升高0.5℃左右,并在黄体期维持在此水平。基础体温的升高与下列哪种激素有关 B A、雌激素 B、孕激素 C、卵泡刺激素 D、黄体生成素 E、甲状腺激素

组织学男性生殖系统习题

17 男性生殖系统 一、选择题 (一)A型题 1.关于精原细胞的描述下述哪项错误？ A.紧贴生精小管的基膜 B.分A、B 两型 C.A型精原细胞又分Ad型与Ap型 D.染色体组型为44＋XY E.B型精原细胞就是精子发生的干细胞 2.关于初级精母细胞的描述错误的就是 A.位于精原细胞近腔侧 B.细胞大,核大而圆,染色质粗网状 C.染色体组型为46, XY D.经两次成熟分裂形成4个次级精母细胞 E.成熟分裂过程历时较长,故生精小管切面上易于见到 3.关于次级精母细胞的描述错误的就是 A.位置靠近管腔 B.较初级精母细胞小,核圆,染色较深 C.染色体核型为23, X或23,Y D.经DNA复制与第2次成熟分裂后形成精子细胞 E.存在时间短,故生精小管切面上不易见到 4.生精细胞第一次减数分裂发生于下述哪一过程？ A.精原细胞变为初级精母细胞 B.初级精母细胞变为次级精母细胞 C.次级精母细胞变为精子细胞 D.精子细胞变为精子 E.精原细胞自身增殖 5.生精小管内最先形成的单倍体细胞就是 A.精原细胞 B.初级精母细胞 C.次级精母细胞 D.精子细胞 E.精子 6.青春期以前的生精小管中一般只有 A.支持细胞 B.精原细胞 C.初级精母细胞 D.支持细胞与精原细胞 E.支持细胞与初级精母细胞 7.具有44＋XY染色体与4n DNA的生精细胞就是 A.精原细胞 B.初级精母细胞 C.次级精母细胞 D.精子细胞

E.Leydig细胞 8.具有22＋X或22＋Y染色体与2n DNA的生精细胞就是 A.精原细胞 B.初级精母细胞 C.次级精母细胞 D.精子细胞 E.精子 9.生精上皮内的生精干细胞就是 A.Ad型精原细胞 B.Ap型精原细胞 C.B型精原细胞 D.初级精母细胞 E.次级精母细胞 10.下述哪一过程既不发生有丝分裂也不发生减数分裂？ A.精原细胞变为初级精母细胞 B、初级精母细胞变为次级精母细胞 C.次级精母细胞变为精子细胞 D.精子细胞变为精子 E.间质细胞的增殖 11.有关精子细胞的描述错误的就是 A.位于生精上皮近腔室 B.分圆型精子细胞与长型精子细胞两类 C.就是单倍体细胞 D.细胞小,核,染色质致密 E.每个细胞经变态过程形成2条精子 12.有关人类精子形态结构的描述错误的就是 A.形似蝌蚪 B.分头、尾两部 C.头部有浓缩的细胞核、顶体与1个中心粒 D.尾部分为颈段、中段、主段与末段 E.中段包有外周致密纤维与线粒体鞘 13.精子的轴丝来自于 A.线粒体 B.高尔基复合体 C.核糖体 D.中心粒 E.滑面内质网 14.由一个精原细胞形成的各级生精细胞同步发育的基础就是 A.细胞间有细胞质桥相连 B.细胞间有紧密连接 C.细胞间有缝隙连接 D.由同一个支持细胞提供营养 E.以上都正确 15.有关睾丸支持细胞的描述下述哪项错误?

组织学女性生殖系统习题

18女性生殖系统一、选择题 （一）A型题 1．关于原始卵泡的叙述错误的是 A．由初级卵母细胞和单层立方的卵泡细胞构成B．卵母细胞的核大而圆，着色浅，核仁明显C．卵母细胞可长期停留于第1次减数分裂的前期D．数量多，位于皮质浅层 E．卵细胞由胚胎期的卵黄囊迁移分化而来 2．卵巢的白体是 A．排卵后组织修复而成 B．排卵后的卵泡壁增生形成 C．卵泡闭锁后膜细胞增生形成 D．黄体退化被结缔组织取代而成 E．间质腺退化被结缔组织取代而成 3．月经后使子宫内膜上皮修复的主要是 A．残留的子宫内膜上皮细胞 B．基底层残留的子宫腺细胞 C．螺旋动脉的内皮细胞 D．基质细胞 E．成纤维细胞

4．关于闭锁卵泡叙述错误的是 A．卵母细胞增大 B．透明带塌陷，不规则 C．卵泡壁坍塌 D．白细胞浸润 E．卵泡膜的膜细胞增大 5．关于黄体描述错误的是 A．新鲜时呈黄色 B．毛细血管极少 C．粒黄体细胞大，数量多，染色浅 D．膜黄体细胞小，数量少，染色深 E．黄体细胞具有分泌类固醇激素细胞的特征6．输卵管黏膜皱襞最发达的部位是 A．漏斗部 B．壶腹部 C．子宫部 D．峡部 E．子宫部和峡部 7．关于子宫颈的结构和功能特点错误的是A．黏膜上皮为单层柱状 B．外口处上皮移行为变移上皮

C．上皮移行处是宫颈癌的好发部位 D．黏膜的分泌受卵巢激素的影响 E．妊娠时分泌物黏稠，可阻止微生物进入子宫8．有关阴道描述错误的是 A．上皮较厚，属角化的复层扁平上皮 B．上皮细胞内聚集大量糖原 C．浅层上皮细胞可脱落 D．脱落细胞的糖原转变为乳酸，使阴道液呈酸性E．上皮的脱落和新生与卵巢活动周期有关 9．卵巢的间质腺是 A．原始卵泡闭锁时由卵母细胞形成的 B．初级卵泡闭锁时由卵泡细胞形成的 C．次级卵泡闭锁时由颗粒细胞形成的 D．次级卵泡闭锁时由膜细胞形成的 E．由基质细胞分化形成的 10．在妊娠后期，乳汁内的初乳小体是 A．乳蛋白凝块 B．吞噬脂滴的巨噬细胞 C．分泌抗体的浆细胞 D．脱落的导管上皮细胞 E．脱落的腺细胞

生理学复习题——12生殖

第十二章生殖 一、单项选择题 1．下列不属于女性副性征的是( ) A．乳腺发达B．喉结突出C．骨盆宽阔 D．皮下脂肪丰满E．音调高 2．关于睾丸的生精功能、叙述正确的是( ) A．支持细胞生成精子B．原始的生精细胞为精原细胞 C．到老年期无生精功能D．精子生成的适宜温度37℃ E．X线照射可促进生精作用 3．促进蛋白质合成和骨髓造血的激素是( ) A．糖皮质激素B．雌二醇C．孕激素 D．雄激素E．醛固酮 4．结扎输精管后将导致( ) A．无精子产生，副性征消失B．有精子产生、副性征消失C．精子排出受阻，性功能受影响D．精子排出受阻、性功能不受影响E．精子排出受阻，副性征消失 5．关于卵巢的生卵功能的叙述，错误的是( ) A．成年女性卵巢中有数万个初级卵泡B．FSH促进卵泡生长发育成熟C．育龄妇女通常每月有一个卵泡成熟D．卵子受精后黄体继续发育E．排卵后的残存卵泡即无功能 6．卵泡刺激素可促进( ) A．卵泡发育成熟B．排卵和黄体形成 C．子宫内膜增生分泌D．睾丸分泌雄激素 E．子宫内膜脱落形成月经 7．关于雌激素的作用，叙述错误的是( ) A．使子宫内膜增生，腺体分泌B．激发并维持女性副性征 C．提高子宫肌对催产素的敏感性D．促进保H2O、保Na+ E．促进阴道上皮增生，角化并合成糖原 8．关于月经的下述说明中，错误的是( ) A．育龄妇女每月一次子宫内膜脱落出血、阴道流血 B．青春期前下丘脑、腺垂体未发育成熟无月经 C．女子12~14岁开始，首次月经，称初潮 D．45~50岁妇女由于子宫萎缩，进入绝经期

E．月经周期受下丘脑—腺垂体—性腺轴调节 9．育龄妇女排卵时间在( ) A．分泌期中间B．基础体温最高时C．月经期中间D．增生期末E．月经停止后第14天前后 10．形成子宫内膜崩溃、脱落，出现月经的直接原因是( ) A．腺垂体分泌促性腺激素减少B．雌激素和孕激素分泌迅速增加C．雌激素和孕激素分泌迅速减少D．雌激素分泌增加，孕激素分泌减少E．雌激素分泌减少，孕激素分泌增加 11．正常月经周期中出现黄体生成素分泌高峰是由于( ) A．促卵泡激素的作用B．雌激素的正反馈作用 C．雌激素的负反馈作用D．孕激素的正反馈作用 E．孕激素的负反馈作用 12．引起卵巢排卵的原因主要是血中呈现( ) A．雌激素高峰B．孕激素高峰C．卵泡刺激素高峰D．黄体生成素高峰E．雌激素、孕激素降低 13．保证胚泡着床，维持妊娠的激素是( ) A．雌激素B．孕激素C．催乳素D．催产素E．生长素14．结扎输卵管后的妇女( ) A．不排卵、有月经B．不排卵，无月经 C．仍排卵、无月经D．仍排卵、有月经、能受精 E．仍排卵、有月经、不能受精 15．妊娠后期血中高浓度的雌激素和孕激素来自( ) A．肾上腺皮质B．胎盘C．卵巢D．卵泡E．妊娠黄体二、多项选择题 1．睾酮的作用是( ) A．刺激男性附性器官生长发育并出现副性征 B．促进蛋白质合成和骨骼肌生长 C．促进下丘脑分泌促性腺激素 D．刺激曲细精管产生精子，维持性欲 E．增强骨髓造血功能 2．雌激素的作用有( ) A．促使子宫平滑肌增厚B．降低子宫肌对催产素的敏感性 C．刺激乳腺腺泡的发育D．促进子宫内膜增生 E．促使阴道上皮细胞增生、角化、合成糖原

组织学女性生殖系统习题

18 女性生殖系统 一、选择题 （一）A型题 1．关于原始卵泡的叙述错误的是 A．由初级卵母细胞和单层立方的卵泡细胞构成B．卵母细胞的核大而圆，着色浅，核仁明显C．卵母细胞可长期停留于第1次减数分裂的前期D．数量多，位于皮质浅层 E．卵细胞由胚胎期的卵黄囊迁移分化而来 2．卵巢的白体是 A．排卵后组织修复而成 B．排卵后的卵泡壁增生形成 C．卵泡闭锁后膜细胞增生形成 D．黄体退化被结缔组织取代而成 E．间质腺退化被结缔组织取代而成 3．月经后使子宫内膜上皮修复的主要是 A．残留的子宫内膜上皮细胞 B．基底层残留的子宫腺细胞 C．螺旋动脉的内皮细胞 D．基质细胞 E．成纤维细胞 4．关于闭锁卵泡叙述错误的是 A．卵母细胞增大 B．透明带塌陷，不规则 C．卵泡壁坍塌 D．白细胞浸润 E．卵泡膜的膜细胞增大 5．关于黄体描述错误的是 A．新鲜时呈黄色 B．毛细血管极少 C．粒黄体细胞大，数量多，染色浅 D．膜黄体细胞小，数量少，染色深 E．黄体细胞具有分泌类固醇激素细胞的特征6．输卵管黏膜皱襞最发达的部位是 A．漏斗部 B．壶腹部 C．子宫部 D．峡部 E．子宫部和峡部 7．关于子宫颈的结构和功能特点错误的是A．黏膜上皮为单层柱状 B．外口处上皮移行为变移上皮

C．上皮移行处是宫颈癌的好发部位 D．黏膜的分泌受卵巢激素的影响 E．妊娠时分泌物黏稠，可阻止微生物进入子宫8．有关阴道描述错误的是 A．上皮较厚，属角化的复层扁平上皮 B．上皮细胞内聚集大量糖原 C．浅层上皮细胞可脱落 D．脱落细胞的糖原转变为乳酸，使阴道液呈酸性E．上皮的脱落和新生与卵巢活动周期有关 9．卵巢的间质腺是 A．原始卵泡闭锁时由卵母细胞形成的 B．初级卵泡闭锁时由卵泡细胞形成的 C．次级卵泡闭锁时由颗粒细胞形成的 D．次级卵泡闭锁时由膜细胞形成的 E．由基质细胞分化形成的 10．在妊娠后期，乳汁内的初乳小体是 A．乳蛋白凝块 B．吞噬脂滴的巨噬细胞 C．分泌抗体的浆细胞 D．脱落的导管上皮细胞 E．脱落的腺细胞 （三）X型题 31．卵泡膜 A．由卵泡周围的结缔组织分化而来 B．可区分出内、外两层 C．外层含卵泡细胞较多 D．内层含膜细胞较多 E．膜细胞有内分泌功能 32．在青春期后卵泡的生长发育过程中 A．卵泡细胞由扁平变为立方、柱状，并增殖成多层B．由初级卵母细胞逐渐发育为成熟卵细胞 C．卵母细胞与卵泡细胞间形成透明带 D．卵泡细胞增殖分化为卵泡膜 E．卵母细胞与卵泡细胞间的缝隙连接使二者协调发育33．妊娠黄体 A．在胎盘分泌的HCG作用下发育增大 B．一般存在14天左右 C．是一个实心细胞团 D．粒黄体细胞较膜黄体细胞数量多 E．分泌孕酮、雌激素和松弛素 34．出现月经期变化的原因是 A．卵巢内黄体退化 B．卵巢内卵泡发育

《生理学》各章知识点总结

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要分泌糖皮质激素（皮质醇）和少量性激素。 5.（1993）性腺可分泌的激素有：A 答案：A层次：记忆考点：性腺的生理功能 解析：性腺的生理功能包括生成生殖细胞和合成分泌性激素。 二、支持细胞： 抑制素（抑制卵泡刺激素的合成与分泌） 转运蛋白：雄激素结合蛋白、转铁蛋白、细胞内视黄醛结合蛋白等 A型题 6.（2017）分泌雄激素结合蛋白的细胞是： A 睾丸精细胞 B 睾丸间质细胞 C 睾丸支持细胞 D 睾丸生精细胞答案：C层次：记忆考点：支持细胞的功能 解析：睾丸支持细胞可合成一些转运蛋白，例如雄激素结合蛋白、转铁蛋白、细胞内视黄醛结合蛋白等。 知识点2：雄激素 一、生物学作用 影响胚胎分化 维持生精作用 促进附性器官的生长、维持性欲 对代谢的影响：促进蛋白质合成 A型题 7.（1990）雄激素的作用，哪项是错误的： A 刺激雄性附性器官发育，并使之维持在成熟状态 B 刺激男性副性征出现 C 促进肌肉与骨骼生长，使男子身高在青春期冲刺式生长 D 分泌过盛，可使男子身高超出常人 E 维持正常的性欲 答案：D层次：理解考点：雄激素的生物学作用 解析：雄激素会导致骨骺与长骨的融合，因此，雄激素分泌过多，会导致骨骺

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组织学与胚胎学生殖系统作业

2011-2012（2）生殖系统试卷 (卷面共有43题,总分100分,各大题标有题量和总分,每小题标号后有小分)一、A选择题(17小题,共17分) [1分](1)睾丸纵隔由（） A、白膜在睾丸后缘增厚形成 B、鞘膜在睾丸后缘增厚形成 C、疏松结缔组织在睾丸中间增厚形成 D、白膜在睾丸前缘增厚形成 E、鞘膜在睾丸前缘增厚形成 [1分](2)生精小管内第二次减数分裂产生的细胞是（） A、精子细胞 B、次级精母细胞 C、精原细胞 D、初级精母细胞 E、精子[1分](3)生精小管内进行第二次减数分裂的细胞是（） A、精子细胞 B、次级精母细胞 C、精原细胞 D、初级精母细胞 E、精子[1分](4)下列对精子发生的论述，错误的是（） A、是精子细胞转变成精子的过程 B、是从精原细胞到形成精子的过程 C、是精原细胞的增殖过程 D、是精母细胞的减数分裂过程 E、是从精母细胞到形成精子的过程 [1分](5)在生精上皮中最不易见到的细胞是（） A、精子细胞 B、精子 C、初级精母细胞 D、次级精母细胞 E、支持细胞[1分](6)下列对精子顶体的描述，正确的是（） A、覆盖核的后1/3 B、内含多种水解酶 C、位于精子的头部和尾部之间 D、可产生雄激素结合蛋白 E、由滑面内质网演变而成 [1分](7)构成血睾屏障的主要结构是（）

A、支持细胞间的紧密连接 B、支持细胞与肌样细胞之间的紧密连接 C、支持细胞间的缝隙连接 D、支持细胞与肌样细胞之间的缝隙连接 E、支持细胞与精原细胞之间的紧密连接 [1分](8)精子细胞的结构特点是（） A、直径约8m,核圆，染色质粗大 B、直径约12m,核椭园，染色质粗大 C、直径约12m,核椭圆，染色质细密 D、直径约8m;核椭园，染色质细密 E、直径约8m,核圆，染色质细密 [1分](9)支持细胞内相对欠发达的细胞器是（） A、滑面内质网 B、粗面内质网 C、高尔基复合体 D、线粒体 E、溶酶体 [1分](10)男性体内雄激素可来自（） A、肾上腺皮质网状带细胞、垂体细胞 B、肾上腺皮质束状带细胞、睾丸间质细胞 C、睾丸间质细胞、甲状腺滤泡旁细胞 D、睾丸间质细胞、肾上腺皮质网状带细胞 E、以上都不对 [1分](11)妊娠黄体能维持（） A、3~4个月 B、4~6个月 C、7~8个月 D、8~9个月 E、9~10个月[1分](12)输卵管上皮分泌细胞功能最活跃时，子宫内膜处于（） A、月经前期 B、月经后期 C、分泌后期 D、排卵前后 E、增生前期[1分](13)月经期后，修复子宫内膜上皮的是（） A、基质细胞 B、螺旋动脉内皮细胞 C、残留的子宫腺上皮细胞