MALE REPRODUCTIVE SYSTEM Reproductive System To Accompany: Anatomy and Physiology Text and Laboratory Workbook, Stephen G. Davenport, Copyright 2006, All Rights Reserved, no part of this publication can be used for any commercial purpose. Permission requests should be addressed to Stephen G. Davenport, Link Publishing, P.O. Box 15562, San Antonio, TX, 78212 Accessory Sex Organs Testes • The primary sex organs of the male are the paired testes. • The testes produce sex hormones and gametes called spermatozoa (sperm). Two of the hormones produced by the testes are – (1) the male sex hormone, testosterone, and – (2) the hormone, inhibin, which regulates the production of sperm (spermatogenesis). • The accessory sex organs include a system of ducts and glands which contribute secretions in the production of semen, the combination of sperm and fluid. • Starting with each testis, the sperm follow a sequence of ducts: – – – – (1) the epididymis, (2) the ductus deferens (vas deferens), (3) the ejaculatory duct, and the (4) urethra, which opens to the outside at the tip of the penis. Accessory Sex Organs • The glands which contribute secretions to the sperm (production of semen) during ejaculation are – (1) the paired seminal vesicles (enter the ejaculatory duct), – (2) the prostate (enters the prostatic urethra), and – (3) the paired bulbourethral glands (enter the spongy urethra). • The external genitalia include – (1) the penis and – (2) the scrotum. Figure 28.1 Male reproductive system in sagittal section. 1 Male Reproductive System • Scrotum – The scrotum is the pouch that contains the testes. The scrotum is divided into halves by a midline septum for the housing of the right and left testes. • Testes – The paired testes produce sperm and hormones. – Seminiferous tubules are located within the testes and are the sites of the production of sperm and the hormone, inhibin, which is involved in the regulation of sperm production. – Located in the connective tissue surrounding the seminiferous tubules are the interstitial (Leydig) cells which produce the sex hormone, testosterone. Male Reproductive System • Seminal vesicles – The paired seminal vesicles are located posterior to the bladder. The alkaline fluid produced by each seminal vesicle is transported to an ejaculatory duct by each seminal vesicle’s duct. During ejaculation, seminal fluid and sperm mix in the ejaculatory duct and are propelled into the urethra (prostatic). • Urethra – The urethra is the tube that serves both the urinary and reproductive systems and is divided into three regions, – (1) the prostatic urethra (passes through the prostate gland) – (2) the membranous urethra (passes through the urogenital diaphragm) – (3) the spongy urethra (passes through the penis and opens to the exterior) Male Reproductive System • Epididymis (pl. epididymides) – An epididymis is located posterior to each testis. The epididymis receives sperm from the testis and is the site where the sperm are stored and develop motility. During ejaculation, smooth muscle contractions within the walls of its tubules force sperm into the ductus (vas) deferens. • Ductus (vas) deferens (pl. vasa deferentia) – The ductus deferens is the tube that transports sperm from the epididymis to the ejaculatory duct. The paired ejaculatory ducts, are located within the prostate. Each is formed by the union of a ductus deferens and a duct from a seminal vesicle. An ejaculatory duct propels sperm (from the vas deferens) and the secretions from a seminal vesicle into the urethra (prostatic). Male Reproductive System • Prostate gland – The prostate gland is located inferior to the bladder and surrounds the urethra (prostatic). – It produces an alkaline fluid that during ejaculation enters the urethra (prostatic) through several small openings. • Bulbourethral glands – The paired bulbourethral glands are located inferior to the prostate. They secrete an alkaline fluid that enters the spongy (penile) urethra. Male Reproductive System • Penis – The penis is attached by its proximal portion called the root. The root leads to the body (shaft) that terminates in the glans penis. – The glans penis is surrounded by the prepuce (foreskin) unless the prepuce was removed by a procedure called circumcision. – The body of the penis consists of three regions of erectile tissue, two dorsal corpora cavernosa (only one corpus cavernosum is seen in a sagittal section) and a medial corpus spongiosum. The corpus spongiosum surrounds the spongy (penile) urethra. – The glans penis surrounds the terminus of the urethra, the external urethral orifice. TESTES The testes are located in the scrotum. They are slightly oval shaped and average about two inches in length and one inch in width. 2 Testes Lab Activity 1 - Testes • Each testis is surrounded by two layers of connective tissues, the – (1) tunica vaginalis- forms a capsule (lined with serous membranes) which surrounds the testis – (2) tunica albuginea- lines the surface of the testis. A portion of the tunica albuginea folds into the testis and divides it into many compartments called lobules. Each lobule consists of one to three coiled seminiferous tubules, which contain the germinal (spermatogenic) epithelium. Lab Activity 1 - Testes Figure 28.3 Scanning power photograph of a cross section of a testis (rat). The testis houses numerous highly convoluted seminiferous tubules and interstitial cells. Lab Activity 1 - Testes • Tunica albuginea – The tunica albuginea is the connective tissue layer covering the surface of the testis. • Interstitial (Leydig) cells – The interstitial (Leydig) cells are located in the connective tissue around the seminiferous tubules. The interstitial (Leydig) cells are endocrine in function and produce the hormone testosterone. Figure 28.2 Scanning power photograph of a section of a testis and its associated epididymis. The testis houses numerous highly convoluted seminiferous tubules. Lab Activity 1 - Testes Figure 28.4 Scanning power photograph of a portion of a longitudinal section of a testis (human) showing its organization into lobules. Each lobule contains one to three highly convoluted seminiferous tubules Lab Activity 1 - Testes • Seminiferous tubules – The seminiferous tubules are the convoluted tubules within the testis where sperm production occurs by the process called spermatogenesis. – Additionally, the seminiferous tubules are the sites of the production of the regulatory hormone called inhibin. The tubules are associated with connective tissues, lymphatic vessels, blood vessels (capillaries are especially abundant), and interstitial (Leydig) cells. 3 Lab Activity 1 - Testes • Spermatogenesis – Spermatogenesis is the process of the production of the male gametes, the spermatozoa, or sperm. – Spermatogenesis involves • the meiotic division of primary spermatocytes to secondary spermatocytes (meiosis I), • the meiotic division of the secondary spermatocytes to spermatids (meiosis II), and the • transformation of the spermatids into spermatozoa (spermiogenesis). • Spermiogenesis – Spermiogenesis is the process of transformation of spermatids into spermatozoa. MEIOSIS and Gamete Production Meiosis is nuclear division in gamete producing cells that results in a reduction in the number of chromosomes from diploid (2n) to haploid (n). Meiosis Meiosis • In meiosis, one of each pair of homologous chromosomes is distributed to a daughter cell. • Haploid daughter cells function as gametes, sperm and eggs. • The number of chromosomes that are found in the somatic (body) cells of the organism is referred to as the diploid number (2n). The diploid number of chromosomes in humans is 46. • Each chromosome has a homologous chromosome, a chromosome that carries the same genes (mostly). Thus, in humans there are 23 pairs of homologous chromosomes. • The set of one each of the homologous chromosomes, or the set of different chromosomes, is described as the haploid (n) number. • The haploid chromosome number for the human is 23 chromosomes, and it is this number found in the sperm and egg. • Before meiosis begins, the chromosomes are replicated • Meiosis is divided into meiosis I and meiosis II Meiosis I Meiosis I • Meiosis I – In meiosis I, the replicated homologous chromosomes are paired, an event called synapsis. – During synapsis crossing-over of portions of the chromosomes (chromatids) occurs. – After crossing-over occurs, the pairs are separated, and one replicated chromosome (a homologous chromosome) is distributed to each daughter cell. • Thus, the daughter cells (n) of Meiosis I have only one of each type of replicated chromosome. Figure 28.5 4 Meiosis II Meiosis II • Entering into meiosis II are the daughter cells from meiosis I. • These cells have one of each type of chromosome (one homolog) in the replicated form (a replicated chromosomes consists of two sister chromatids). • In meiosis II, the sister chromatids are separated and one of each sister chromosome is distributed to the daughter cells. Thus, the daughter cells, which function as gametes, have only one of each homologous chromosome. • The gametes are haploid (n). Fusion of the gametes, a sperm and an egg, during fertilization produces the diploid (2n) number of chromosomes and produces a single cell called a zygote. Figure 28.5 Meiosis - Spermatogenesis • Spermatogenesis is the process of male gamete (sperm) production. Spermatogenesis occurs in the seminiferous tubules of the testis. • Precursor (stem) cells called spermatogonia, located at the periphery of the seminiferous tubules, undergo mitotic divisions. Some of the daughter cells remain at the periphery of the tubules as precursor cells. Others differentiate into primary spermatocytes (2n.) • Primary spermatocytes enter meiosis I and produce two haploid daughter cells called secondary spermatocytes (n.) • The two secondary spermatocytes enter meiosis II and each produces two haploid spermatids (n.) • Spermatids undergo a process of transformation called spermiogenesis and develop into spermatozoa (sperm.) Meiosis - Spermatogenesis Figure 28.6 Spermatogenesis is the process of male gamete production. Meiosis reduces the diploid human chromosome number of 46 to the haploid number of 23. Meiosis assures that each of the gametes contains only half of the homologous chromosomes. Lab Activity 2 – Testis Lab Activity 2 – Testis Spermatogenesis - Microscopic Study Spermatogenesis - Microscopic Study Figure 28.7 Section of testis showing seminiferous tubules, c.s. Figure 28.8 High power photograph of a seminiferous tubule (rat) showing cell division. Dividing cells are not always observed in testis preparations. 5 Lab Activity 2 – Testis Lab Activity 2 – Testis Spermatogenesis - Microscopic Study Spermatogenesis - Microscopic Study Figure 28.9 Low power photograph of a seminiferous tubule. The specific identification of the cell types is difficult and is based mostly by their sequence of development. Figure 28.10 High power photograph of a seminiferous tubule (rat) showing sustentacular cells and spermatogenic cells. The cells are mostly identified by the location and shape of their nuclei. Interstitial cells are located around the seminiferous tubule. Sustentacular cells Sustentacular cells • The sustentacular cells are permanent residents of the germinal epithelium and undergo very little if any cell division. • The sustentacular cells occupy the space between the outer margin of the seminiferous tubule and its lumen. The sustentacular cells partially invest the spermatogenic cells, which causes the spermatogenic cells to be located between the plasma membranes of adjacent sustentacular cells. • Sustentacular cells function to support the cells of spermatogenesis, and thus, are often called “nurse” cells. • In addition to metabolically supporting the spermatogenic cells, sustentacular cells function to support spermatogenesis by the production of the blood-testis barrier. • The blood-testis barrier is necessary to prevent an auto-immune response toward the haploid spermatogenic cells located toward the lumen of the seminiferous tubule. • The fusion of the membranes of the adjacent sustentacular cells divides the seminiferous tubes into an outer basal compartment and an inner luminal compartment. Sustentacular cells Sustentacular cells Figure 28.11 Sustentacular cells fuse by tight junctions to form a blood-testis barrier. The sustentacular cells fuse to divide the seminiferous tubule into a basal compartment and a luminal compartment. The inner luminal compartment, which contains haploid cells, has no direct blood exchange. Figure 28.11 Sustentacular cells fuse by tight junctions to form a blood-testis barrier. The sustentacular cells fuse to divide the seminiferous tubule into a basal compartment and a luminal compartment. The inner luminal compartment, which contains haploid cells, has no direct blood exchange. 6 Follicle Stimulating Hormone (FSH) MALE HORMONAL REGULATION Follicle-stimulating hormone (FSH) Follicle Stimulating Hormone (FSH) • Sustentacular Cells • The sustentacular cells also function in the regulation of spermatogenesis by responding to blood levels of follicle stimulating hormone (FSH). • FSH promotes the release of androgen binding protein (ABP) from the sustentacular cells. – Androgen binding protein targets the spermatogenic cells and results in their increased uptake of testosterone. Increased stimulation of the spermatogenic cells by testosterone results in an increased rate of spermatogenesis Follicle Stimulating Hormone (FSH) • Sustentacular Cells • Sustentacular cells also release the hormone inhibin, which functions in the regulation of the release of follicle stimulating hormone (FSH) from the anterior pituitary gland. – Increased stimulation of the sustentacular cells by FSH, increases inhibin release, which by negative feedback targets the hypothalamus, reduces the release of GnRH, which reduces the release of FSH. – Decreased levels of inhibin result in hypothalamic increase of gonadotropin-releasing hormone (GnRH), which results in an increase of follicle-stimulating hormone (FSH). Luteinizing hormone (LH) • Luteinizing hormone (LH) is produced and released from the anterior pituitary gland (pars distalis). • Luteinizing hormone (LH) targets the interstitial cells of the testes and increases the release of testosterone. • Negative feedback controls the secretion of luteinizing hormone (LH). Increased levels of testosterone result in hypothalamic reduction of gonadotropin-releasing hormone (GnRH). Figure 28.12 Follicle-stimulating hormone (FSH) is produced and released from the anterior pituitary (pars distalis). Follicle-stimulating hormone (FSH) targets the sustentacular cells. Sustentacular cells release androgen binding protein (ABP) and inhibin. Androgen binding protein targets the spermatogenic cells and inhibin targets the hypothalamus. Luteinizing Hormone (LH) Figure 28.13 The release of luteinizing hormone from the anterior pituitary is regulated by gonadotropin-releasing hormone (GnRH). Luteinizing hormone (LH) is produced and released from the anterior pituitary (pars distalis). Luteinizing hormone (LH) targets the interstitial cells to increase testosterone production. 7 Spermatozoa (Sperm) SPERMATOZOA (SPERM) Spermatozoa are the gametes of the male. Spermatozoa (Sperm) • Head of Sperm – The head of the spermatozoon contains the nucleus of the cell. – A structure called the acrosomal cap (or acrosome) is located at the tip of the head. The acrosomal cap releases enzymes which function in the penetration of the egg. • The nucleus of each sperm contains one of each homologous chromosome for a total of 23 chromosomes. • A spermatozoon is divided into three regions, a – head, – midpiece, and the – tail. Spermatozoa (Sperm) • Midpiece of Sperm – The midpiece of the spermatozoon is connected to the head by a short neck. – The neck is the origin of the microtubules which pass through the center of the midpiece into the tail, or flagellum. Numerous mitochondria are located at the periphery of the microtubules and provide the energy (ATP) to drive the movement the microtubules. – The movement of the microtubules results in the movement of the tail (flagellum). Simple sugars such as fructose are absorbed from the fluid surrounding the spermatozoa and serve as the primary fuel molecule in the production of ATP. • Tail of Sperm – The tail, or flagellum, contains microtubules and functions to move the spermatozoon. Lab Activity 3 - Sperm EPIDIDYMIS The ductus epididymis functions as the site where sperm are stored and mature. Figure 28.14 Oil immersion photograph of human spermatozoa. A spermatozoon is divided into a head, midpiece, and a tail (flagellum). 8 Epididymis Epididymis • \The seminiferous tubules of the testis merge into a region of the testis called the rete testis. The rete testis consists of a group of small ducts lined with simple cuboidal epithelium that deliver spermatozoa into the efferent ducts located in the head of the epididymis. The efferent ducts (ductuli efferentes) are lined with ciliated columnar epithelium and merge to form the highly coiled long duct of the body and tail of the epididymis, the ductus epididymis. • The ductus epididymis functions as the site where sperm are stored and mature. The ductus epididymis forms the ductus deferens, which connects the epididymis to the ejaculatory duct. • The short ejaculatory duct passes through the prostate gland and enters the urethra. • The ductus epididymis consists of pseudostratified columnar epithelium with long stereocilia (microvilli). • The stereocilia do not function in movement, but function to increase surface area for the secretion of glycoproteins and the absorption of fluid released by the testis. • The glycoproteins function in the process of spermatozoa maturation; the spermatozoa become physiologically capable of fertilization of the egg. Lab Activity 4 - Epididymis Lab Activity 4 - Epididymis Figure 28.15 Scanning power photograph of a section of the epididymis and its associated testis. The body and tail of the epididymis contains the ductus epididymis, a long highly convoluted tube. Figure 28.16 Low power photograph of a section of the epididymis. The body and tail of the epididymis consists of a long highly coiled tube, the ductus epididymis. Lab Activity 4 - Epididymis DUCTUS DEFERENS The ductus deferens, or vas deferens, originates from the ductus epididymis at the tail of the epididymis. Figure 28.17 High power photograph of a section of the ductus epididymis. Stereocilia (microvilli) extend from the columnar epithelial cells and function to increase the surface area for secretion of glycoproteins and absorption of fluid. 9 Ductus (vas) Deferens • The ductus deferens, or vas deferens, originates from the ductus epididymis at the tail of the epididymis. It turns upward to leave the scrotum as a component of the spermatic cord and enters the abdominal cavity at the opening of the inguinal canal (ring). • From here the ductus deferens enters the pelvic cavity ultimately to pass medially to the seminal vesicle and downward to the base of the prostate gland. • At the base of the prostate gland the ductus deferens merges with the duct from the seminal vesicle to form the ejaculatory duct, which passes through the prostate gland and enters the prostatic urethra. • The ductus deferens functions to transport sperm from the epididymis to the ejaculatory duct. Movement of sperm during ejaculation is by peristaltic waves of the muscularis. Lab Activity 5 – Ductus (vas) Deferens Figure 28.18 Low power photograph of a section of the ductus deferens. The ductus deferens is a muscular tube with a small lumen. Its muscular layer consists of smooth muscle which functions in the movement of sperm by peristalsis. Seminal Vesicles SEMINAL VESICLES The two seminal vesicles are located posterior to the urinary bladder and anterior to the rectum Lab Activity 6 – Seminal Vesicles Figure 28.19 Low power photograph of a section of the seminal vesicle (primate). The seminal vesicle consists of a highly coiled duct. • The two seminal vesicles are located posterior to the urinary bladder and anterior to the rectum. Each is associated with its respective ductus deferens by way of an ejaculatory duct formed from the union of the ductus deferens and the duct from its associated seminal vesicle. • A seminal vesicle is formed from a long coiled and highly branched tube. The posterior end of the tube is a blindend, and anteriorly the tube forms a narrow straight vesicular duct that joins with its associated ductus deferens. • The seminal vesicles function to secrete vesicular fluid, the major component (about 60%) of semen. Vesicular fluid is a fructose rich slightly alkaline fluid. Lab Activity 6 – Seminal Vesicles Figure 28.20 Low power photograph of a section of the seminal vesicle (human). This photograph shows a section of the highly coiled duct. The duct has a secretory mucosa that is highly branched (diverticula). 10 Lab Activity 6 – Seminal Vesicles Prostate Gland Figure 28.21 High power photograph of a section of the seminal vesicle (human). This photograph shows a section of the mucosa of the highly coiled duct. The secretory mucosa is highly branched (forms diverticula), which functions to increase its secretory surface area. Prostate Gland • The prostate gland is located inferior to the urinary bladder at the origin of the urethra (prostatic urethra), which it completely surrounds. • Transversing the prostate is the ejaculatory duct, formed from the union of the ductus deferens and the duct from its respective seminal vesicle. The prostate is formed from highly branched glands (tubuloalveolar units), which merge into ducts that enter the prostatic urethra. • The highly branched prostatic glands are lined with columnar epithelium which functions in the secretion of prostatic fluid. Prostatic fluid comprises about 30% of semen and is a slightly acid fluid that mostly functions in the activation of sperm. • The interstitial tissue (tissue between the glands), or stroma, consists mostly of smooth muscle and connective tissue. Contraction of the smooth muscle during ejaculation moves prostatic fluid into the prostatic urethra. Lab Activity 7 – Prostate Gland Figure 28.23 Low power photograph of a section of the prostate (primate). The glands (tubuloalveolar units) are lined with columnar epithelium which functions in the secretion of prostatic fluid. Lab Activity 7 – Prostate Gland Figure 28.22 Scanning power photograph of a section of the prostate (primate). The prostate is composed of glands (tubuloalveolar units) with function in the secretion of prostatic fluid. Lab Activity 7 – Prostate Gland Figure 28.24 High power photograph of a section of the prostate (primate). The glands (tubuloalveolar units) are lined with columnar epithelium which functions in the secretion of prostatic fluid. The glands empty into the prostatic urethra. 11 Penis PENIS The penis is the copulatory organ of the male and is formed from three bodies of erectile, or cavernous, tissue, which are supported by connective tissue and covered with skin. Penis • The three bodies of erectile tissue are the two dorsal corpora cavernosa and the corpus spongiosum. • A portion of the penis is internal and forms the root of the penis. The external portion of the penis is called the shaft. The single ventral corpus spongiosum houses the spongy, or penile, urethra. The corpus spongiosum expands at the tip of the penis to form the glans penis, the head of the penis. • The penis is covered with skin, which folds over the glans penis as the prepuce, or foreskin. The procedure called circumcision removes the prepuce. The three bodies of cavernous tissue are each surrounded by connective tissue called the tunica albuginea. Penis • The tunica albuginea surrounding the two dorsal corpora cavernosa meets in the midline to form the median septum. The tunica albuginea that surrounds the corpus spongiosum is thinner than that surrounding the corpora cavernosa. • The cavernous tissue is formed from large vascular areas (sinuses), or cavernous spaces. Partitions called trabeculae divide the cavernous spaces which fill with blood during erection of the penis. Figure 28.27 The penis is formed from three bodies of erectile, or cavernous tissue that are supported by connective tissue and covered with skin. Lab Activity 8 - Penis Figure 28.28 Scanning power photograph of a section of the human penis. The preparation shows a portion of the corpus cavernosum, corpus spongiosum, and the urethra. Lab Activity 8 - Penis Figure 28.29 Low power photograph of a section of the human penis. The preparation shows a portion of the corpus spongiosum and the urethra. 12 Lab Activity 8 - Penis Lab Activity 8 - Penis Figure 28.30 Low power photograph of a section of the human penis. The preparation shows a portion of the corpus cavernosum. Increase blood flow into the cavernous spaces during sexual arousal increases blood pressure within the spaces resulting in erection. Decreased blood flow allows the elastic and smooth muscle fibers of the trabeculae to recoil, allowing the penis to become flaccid. Figure 28.31 Low power photograph of a section of the human penis. The preparation shows a portion of the corpus cavernosum which has been cleared of blood during processing of the tissue. Female Reproductive System FEMALE REPRODUCTIVE SYSTEM The primary sex organs of the female are the paired ovaries. Female Reproductive System • • • • • • • • • The ovaries produce hormones and gametes called eggs (ova). • Sex hormones produced by the ovaries include (1) estrogen and (2) progesterone. • The accessory sex organs, starting with the ovaries, are the – (1) uterine tubes (Fallopian tubes, or oviducts), – (2) uterus, and – (3) vagina. Female Reproductive System The external genitalia (vulva) include the (1) mons pubis, (2) labia majora, (3) labia minora, (4) vestibule, (5) clitoris (6) vestibular glands, and the (7) paraurethral glands. Figure 28.32 Illustration of a midsagittal section of the female pelvis. 13 External Genitalia • Labia majora – The labia majora are two skin-covered folds of mostly adipose tissue located inferior to the mons pubis and form the lateral boundary of the vulva. EXTERNAL GENITALIA The external genitalia (or vulva) are the external genital organs of the female reproductive system. • Labia minora – The labia minora are two skin-covered folds, each located medially to its respective labium majus. The labia minora form the lateral boundary of the vestibule (opening) of the vagina. • Vestibule – The vestibule is the chamber, or space, that is formed within the boundary of the labia minora. Both the urethra and the vagina open into the vestibule. External Genitalia • Clitoris – The clitoris is a small rounded erectile organ located at the anterior of the vulva at the junction of the labia minora. The prepuce of the clitoris is formed from an extension of the labia minora. • Mons pubis – The mons pubis is an elevated area that covers the pubic bones. After puberty it is covered with hair and is usually soft due to the presence of fat. External Genitalia • Greater vestibular glands (Bartholin’s glands) – There are two greater vestibular glands. One of each vestibular gland is located lateral to the vagina and opens into the vagina’s vestibule. – The glands produce a mucus lubricating fluid during intercourse. The greater vestibular glands are homologous (similar in structure) to the male’s bulbourethral glands. Several smaller glands called lesser vestibular glands also open into the vestibule. • Paraurethral glands – The paraurethral glands open into the external orifice (opening) of the urethra and are homologous to the male’s prostate gland. Internal Organs • Broad ligament INTERNAL ORGANS – The broad ligament is a double fold of the peritoneum that folds over and anchors the ovaries, uterine tubes, and uterus. The broad ligament anchors the uterus to the lateral walls of the pelvis, and between its two layers of peritoneum the broad ligament provides a route for the passage of the uterine tubes. The ovary is attached to the dorsal surface of the broad ligament near the broad ligament’s lateral merging with the peritoneum of the pelvis. • Rectouterine pouch – The rectouterine pouch is the area formed between the uterus and the rectum by the broad ligament. • Vesicouterine pouch – The vesicouterine pouch is the area formed between the uterus and the posterior wall of the bladder by the broad ligament. 14 Internal Organs • Ovary Internal Organs • Ovary – The paired ovaries are located laterally to the uterus and are attached to the dorsal surface of the broad ligament. – Each ovary is supported and anchored by the mesovarium, the ovarian ligament, and the suspensory ligament. – The ovary is lined with modified visceral peritoneum. At the margin of the ovary, the visceral peritoneum changes from simple squamous to simple columnar epithelium, which covers the ovary as the layer of the ovary called the germinal epithelium. The germinal epithelium only functions to structurally line the ovary. – Internally, the ovary is divided into an outer region called the cortex and a deep central region called the medulla. – Cortex The cortex of the ovary is the outer region of the ovary and contains numerous follicles embedded in a surrounding tissue called the stroma. The follicles are the sites of oocyte production and maintenance, and along with the corpus luteum, produce the primary sex hormones of the female, estrogen and progesterone. The follicles also release the regulatory hormone called inhibin. – Medulla The medulla of the ovary is the inner central region of the ovary. The tissue of the medulla, the stroma, provides a pathway for blood vessels, nerves, etc. Internal Organs • Mesovarium – The mesovarium is a sheet of peritoneum that attaches the ovary to the lateral wall of the pelvis. • Ovarian ligament – The ovarian ligament attaches the medial surface of the ovary to the lateral wall of the uterus, at a point just inferior to the entrance of the uterine tube. • Suspensory ligament – The suspensory ligament is a fold of peritoneum that attaches the lateral wall of the ovary to the wall of the pelvis, and serves as a site for the passage of the ovarian blood vessels. Internal Organs • Uterine tubes (Fallopian tubes or Oviducts) – Each of the uterine tubes extends from an ovary to the superior, lateral region of the uterus. – Extending from the uterus toward the ovary is the region called the isthmus. – Near the ovary, they have an expanded region called the ampulla. The open end of the ampulla is called the infundibulum. – Small projections called fimbriae extend from the infundibulum and drape over the ovary. Internal Organs • Uterus – The uterus is located posterior and superior to the bladder. It receives the two uterine tubes and serves as the site for the implantation and development of the fertilized egg. • Three major regions are the – (1) fundus, the rounded portion superior to the entrances of the uterine tubes. – (2) body, the central portion of the uterus – (3) cervix, the narrow portion that projects outward into the vagina. The cervical canal provides a pathway from the vagina to the interior cavity of the uterus. Internal Organs Uterine wall • The three layers of the wall of the uterus are, from the outside inward, the – (1) perimetrium, the serous membrane that covers the outside of the uterus – (2) myometrium, the thick middle smooth muscle layer – (3) endometrium, the inner layer that undergoes cyclic changes in response to ovarian hormones. The endometrium is divided into the stratum functionalis and the stratum basalis. Vagina – The vagina is the tubular organ that extends from the cervix to the exterior region called the vestibule. It serves as a passageway for the delivery of a baby and receives the penis during sexual intercourse. 15 Internal Organs OVARY The paired ovaries are located laterally to the uterus and are attached to the dorsal surface of the broad ligament. Figure 28.33 Anterior view of the uterus and associated organs. Ovary Lab Activity 9 - Ovary • The paired ovaries are located laterally to the uterus and are attached to the dorsal surface of the broad ligament. The ovary is lined with modified visceral peritoneum called the germinal epithelium. • The germinal epithelium functions as a lining of the ovary. Internally, the ovary is divided into an outer region called the cortex and a deep central region called the medulla. – The cortex of the ovary is the outer region of the ovary and contains numerous follicles embedded in a surrounding tissue called the stroma. – The medulla of the ovary is the inner central region of the ovary. The tissue of the medulla, the stroma, provides a pathway for blood vessels, nerves, etc. Lab Activity 9 - Ovary Figure 28.34 Scanning power photograph of an ovary (monkey). Numerous follicles are observed in the cortex. Lab Activity 9 - Ovary Primordial follicles Primordial follicles are located toward the periphery of the cortex. Each primordial follicle consists of a primary oocyte that is surrounded by a single layer of squamous cells called follicular cells. Figure 28.35 Low power photograph of the cortex of an ovary (monkey). Three types of follicles can be observed, primordial, primary, and secondary follicles. Figure 28.36 High power photograph of primordial follicles (monkey ovary). 16 Lab Activity 9 - Ovary Primary follicles Primary follicles develop from primordial follicles. Each primary follicle consists of a primary oocyte that is surrounded by several layers of cuboidal cells called granulosa cells. Figure 28.37 High power photograph of primary follicles (monkey ovary). Secondary follicles (tertiary, vesicular, or Graafian) Secondary follicles (tertiary, vesicular, or Graafian) • Secondary follicles develop from primary follicles. • The secondary follicle consists of an oocyte surrounded by several layers of granulosa cells that produce a fluid-filled central cavity called the antrum. • A layer of granulosa cells, the corona radiata, surrounds the oocyte and the region of glycoproteins, the zona pellucida. The zona pellucida is a glycoprotein matrix with microvilli, microvilli of the granulosa cells and microvilli of the oocyte. The microvilli allow for the exchange of materials between the primary oocyte and the granulosa cells. Lab Activity 9 - Ovary • Upon ovulation, the secondary oocyte and its associated zona pellucida and corona radiata are discharged from the mature secondary follicle. • A mature secondary follicle that is ready to rupture in ovulation is referred to as a vesicular (Graafian) follicle. After ovulation, the vesicular follicle is transformed into an endocrine organ, the corpus luteum. Figure 28.38 High power photograph of an early secondary follicle (monkey ovary). Lab Activity 9 - Ovary CORPUS LUTEUM The corpus luteum develops from the ruptured vesicular (or Graafian) follicle after ovulation Figure 28.39 High power photograph of a late secondary follicle (monkey ovary). Secondary follicles are also called vesicular, tertiary, or Graafian follicles. 17 Corpus Luteum • The corpus luteum develops from the ruptured vesicular (or Graafian) follicle after ovulation. • The corpus luteum functions as an endocrine gland and produces progesterone (mostly) and estrogen. • Hormone producing cells are the theca and granulosa lutein cells. The theca is the outer covering of the Graafian follicle. Some of its cell, the theca lutein cells produce estrogen. The granulosa cells develop into granulosa lutein cells and produce progesterone. • If an embryo implants, a hormone is produced by the developing placenta that maintains the corpus luteum. If an embryo does not develop, the corpus luteum begins to degenerate, and its hormonal functions are terminated by the time of menstruation. The corpus luteum degenerates into the corpus albicans. Lab Activity 10 – Corpus Luteum Figure 28.41 Scanning power photograph of the corpus luteum (monkey ovary). This corpus luteum is relatively late in development as the lutein cells, which develop from the granulosa cells, have completely filled the prior location of the antrum. Lab Activity 10 – Corpus Luteum Figure 28.40 Scanning power photograph of the corpus luteum. This corpus luteum is relatively early in development as the lutein cells, which develop from the granulosa cells, have not completely filled the prior location of the antrum. Lab Activity 10 – Corpus Luteum Figure 28.42 High power photograph of the corpus luteum (monkey ovary). The progesterone-secreting cells of the corpus luteum are the lutein cells which develop from the granulosa cells of the follicles. Lab Activity 11 – Corpus Albicans CORPUS ALBICANS The corpus albicans is the fibrous connective tissue that develops as a result of the degeneration and reabsorption of the corpus luteum. Figure 28.43 Scanning power photograph of the corpus albicans (monkey ovary). The corpus albicans is usually pale-stained as it consists mostly of fibrous connective tissue. 18 Lab Activity 11 – Corpus Albicans MEIOSIS - OOGENESIS Oogenesis is the process of production and maturation of the egg (ovum). Figure 28.44 High power photograph of the corpus albicans (monkey ovary). The pale-stained corpus albicans consists mostly of fibrous connective tissue. Meiosis and Oogenesis • Oogenesis is the process of production and maturation of the egg (ovum). • Oogenesis begins in early fetal development with the mitotic production of oogonia (2n), the precursor (stem) cells to the ova. • The oogonia develop into primary oocytes as they become surrounded by a single layer of follicular cells, forming a primordial follicle. Meiosis and Oogenesis • A primary oocyte (2n) completes meiosis I to produce two cells of very unequal size. The smaller of the two cells is called the first polar body. The larger secondary oocyte (n), which received most of the cytoplasm, continues with meiosis II until metaphase II and then stops. • The secondary oocyte in metaphase II is ovulated. If fertilization occurs, the secondary oocyte finishes meiosis II. It produces a second polar body and an ovum (n), or mature egg. • Nuclear union between the ovum and sperm produces a diploid (2n) zygote. The second polar body, as with the other two second polar bodies produced by meiosis II of the first polar body, degenerates. Meiosis and Oogenesis • The primary oocytes enter meiosis I, which quickly stops in prophase I. At birth and until puberty, the ovary contains several million primary oocytes, each in a primordial follicle. • At puberty and mostly under the influence of gonadotropic hormones (mostly follicle stimulating hormone, FSH) cyclically released from the anterior pituitary, some hormone sensitive primordial follicles begin to develop. The primary oocyte continues with meiosis I, and the single layer of follicular cells develops into layers of cells, the granulosa cells, forming a primary follicle. Meiosis and Oogenesis Figure 28.45 Oogenesis is the process of female gamete production. Meiosis reduces the diploid human chromosome number of 46 to the haploid number of 23. Meiosis assures that the ovum contains only half of the homologous chromosomes. 19 Lab Activity 12 – Oogenesis Figure 28.46 Ovary (human) of a infant. Primary oocytes are produced early in fetal development and their meiotic activity stops in prophase I. The primary oocytes are surrounded by a single layer of follicular cells. A primary oocyte and its follicular cells are called a primordial follicle. Primordial follicles remain inactive until the onset of puberty. Lab Activity 12 – Oogenesis Figure 28.47 Ovary (monkey) showing various stages of oocyte and follicular development. Lab Activity 12 – Oogenesis OVARIAN CYCLE Figure 28.48 Primordial follicles. Each primordial follicle consists of a single layer of follicular cells and a primary oocyte. It is estimated that at birth each ovary contains up to 2,000,000 primordial follicles. The ovarian cycle is the approximate 28 days (monthly) sequence of changes that occur in the ovary in relation to its follicular development, ovulation, and development of the corpus luteum Ovarian Cycle Follicle-stimulating hormone (FSH) The ovarian cycle is the approximate 28 days (monthly) sequence of changes that occur in the ovary in relation to its follicular development, ovulation, and development of the corpus luteum. • The cycle of the ovary is regulated by the two gonadotropic hormones (released from the anterior pituitary (pars distalis) called follicle stimulating hormone (FSH) and luteinizing hormone (LH), and by negative feedback from three hormones, estrogen, progesterone, and inhibin. • Follicle-stimulating hormone (FSH) is produced and released from the anterior pituitary gland (pars distalis). • Follicle-stimulating hormone (FSH) targets the follicles of the ovaries to stimulate follicular development. Developing follicles under the influence of FSH release the regulatory hormone inhibin and mostly the sex hormone estrogen. • 20 Follicle-stimulating hormone (FSH) • A negative feedback mechanism controls the secretion of follicle-stimulating hormone (FSH). – Increased levels of inhibin result in hypothalamic reduction of gonadotropin-releasing hormone (GnRH), which results in a reduction of folliclestimulating hormone (FSH); – decreased levels of inhibin result in hypothalamic increase of gonadotropin-releasing hormone (GnRH), which results in an increase of follicle-stimulating hormone (FSH). Luteinizing Hormone (LH) • Luteinizing hormone (LH) is produced and released from the anterior pituitary (pars distalis). • Luteinizing hormone (LH) targets – (1) the mature follicle of the ovary to trigger ovulation, – (2) and promotes the development of the corpus luteum, which secretes estrogen and progesterone. Follicle-Stimulating Hormone (FSH) Figure 28.49 Follicle-stimulating hormone (FSH) is produced and released from the anterior pituitary gland (pars distalis). Follicle-stimulating hormone (FSH) targets the follicles and promotes their development. Developing follicles release inhibin which results in the reduction of GnRH, thus reducing FSH. Luteinizing Hormone (LH) • A negative feedback mechanism controls the secretion of luteinizing hormone (LH). • Increased levels of estrogen and progesterone result in hypothalamic reduction of gonadotropin-releasing hormone (GnRH), which result in a decrease of luteinizing hormone (LH); • Decreased levels of estrogen and progesterone result in hypothalamic increase of gonadotropinreleasing hormone (GnRH), which results in an increase of luteinizing hormone (LH). Luteinizing Hormone (LH) Figure 28.50 Luteinizing hormone (LH) is produced and released from the pars distalis. For females, luteinizing hormone (LH) targets the mature follicle, and supports corpus luteum and secretion of estrogen and progesterone. Phases of Uterine Cycle The ovarian cycle is divided into the follicular phase and the luteal phase. 21 Phases of Uterine Cycle Phases of Uterine Cycle • Follicular phase – The follicular phase of the ovarian cycle is the time of follicular development, approximately days 1 - 14, with day 14 as the day of ovulation. • Luteal phase – The luteal phase of the ovarian cycle is the time following ovulation, approximately days 14 - 28, when the corpus luteum develops and is maintained. Phases of Uterine Cycle Figure 28.52 Photographs of the structures of an ovarian cycle. During the ovarian cycle follicles develop (follicular phase), ovulation occurs, and the corpus luteum develops (luteal phase). A corpus albicans, a body of fibrous tissue forms from the degeneration of the corpus luteum. Phases of Uterine Cycle Figure 28.54 Photographs of primary and secondary follicles of the ovary (monkey). Primary follicles are developing follicles. Their granulosa and theca cells secrete estrogen. As the number and size of the granulosa and theca cells increases so does the level of estrogen. Primary follicles develop into secondary follicles. Figure 28.51 Illustration showing the sequences of changes in the 28 day ovarian cycle. The follicular phase (days 1 - 14) is the preovulatory phase and is the time of follicular development. The luteal phase (days 14 - 28) is the postovulatory phase and is the time of the development of the corpus luteum. Phases of Uterine Cycle Figure 28.53 Photographs of primordial and primary follicles of the ovary (monkey). Primordial follicles receptive to follicle stimulating hormone (FSH) released by the anterior pituitary gland develop into primary follicles. Phases of Uterine Cycle Figure 28.55 Photographs of an early secondary and a late secondary follicle of the ovary (monkey). Secondary follicles secrete increasing amounts of estrogen as their granulosa and theca cells increase in size and number. A late secondary follicle develops into a vesicular (Graafian) follicle. 22 Phases of Uterine Cycle Figure 28.56 Photograph of an early secondary follicle (monkey ovary) and an illustration of ovulation. At ovulation the follicular phase ends, and the luteal phase begins with the conversion of the vesicular follicle of ovulation into an endocrine gland called the corpus luteum. Phases of Uterine Cycle Figure 28.57 Illustration of a vesicular follicle of ovulation and a photograph of the corpus luteum (monkey ovary). The corpus luteum functions as an endocrine gland and secretes progesterone and estrogen. Phases of Uterine Cycle Figure 28.58 Photographs of the corpus luteum and the corpus albicans (monkey ovary). The corpus albicans consists of fibrous connective tissue that develops from the degeneration of the corpus luteum. Figure 28.59 Illustration of the ovarian cycle. The cycle of the ovary is regulated by the two gonadotropic hormones released from the anterior pituitary (pars distalis), follicle stimulating hormone (FSH) and luteinizing hormone (LH), and by negative feedback from three hormones, estrogen, progesterone, and inhibin. Uterine Tube UTERINE TUBE The uterine tube (also called oviduct, or Fallopian tube) is the tube that extends between the ovary and the uterus • The uterine tube (also called oviduct, or Fallopian tube) is the tube that extends between the ovary and the uterus. • Each of the two uterine tubes is divided into three regions, – (1) isthmus, – (2) ampulla, and – (3) infundibulum. 23 Lab Activity 13 – Uterine Tube Lab Activity 13 – Uterine Tube Figure 28.62 Low power photograph of the ampulla’s wall. The highly folded mucosa extends into the lumen of the uterine tube. Figure 28.61 Scanning power photograph of the isthmus of the uterine tube. Lab Activity 13 – Uterine Tube UTERUS Figure 28.63 High power photograph of the mucosa of the ampulla of the uterine tube. The highly folded mucosa is lined with ciliated columnar cells. Numerous secretory cells are found among the ciliated cells. Uterus The uterus is the organ that functions as the site for the development of the embryo and fetus prior to birth. Uterus • The uterus is located in the pelvic cavity between the urinary bladder and the rectum. • The uterine tubes open into its superior-lateral body. • The cervix of the uterus opens into the inferior vagina. • The uterus is the organ that functions as the site for the development of the embryo and fetus prior to birth. Figure 28.64 The uterus is the organ that functions as the site for the development of the embryo and fetus prior to birth. 24 Uterus Uterus • Isthmus • Fundus – The fundus of the uterus is the superior portion of the uterus, the region above the points of attachment of the uterine tubes. • Body – The body of the uterus is the region between the superior fundus and the inferior isthmus – The isthmus of the uterus is a narrow constricted region inferior to and continuous with the body. The uterine cavity of the uterus narrows to form the internal orifice at the superior internal boundary of the isthmus. • Cervix – The cervix of the uterus is the most inferior portion of the uterus. – The cervix is continuous with the isthmus and its inferior surface curves outward into the vagina. – The opening into the cervix is called the external orifice (or cervical os). The external orifice opens into the cervical canal, which leads to the internal orifice. The internal orifice is continuous with the uterine cavity. Uterine Wall • Perimetrium – The perimetrium is the serous membrane, the peritoneum, which covers the fundus and the anterior-posterior surfaces of the uterus. UTERINE WALL • Myometrium – The myometrium is the thick middle muscularis of the uterus. • Endometrium The wall of the uterus consists of three layers, from the outside inward, the layers are the (1) perimetrium, (2) myometrium, and (3) endometrium. Uterine Wall – The endometrium is the inner mucosa of the uterus and consists of two layers the (1) stratum functionalis and the (2) stratum basalis. The endometrium undergoes cyclic changes, the uterine cycle, in response to ovarian hormones. – The endometrium consists of numerous uterine (endometrial) glands and blood vessels located within a layer of connective tissue, the stroma. Uterine Wall • Uterine glands – Uterine glands are the tubular glands of the mucosa (endometrium) of the uterus. Early in the uterine cycle the uterine glands are small and relative straight. Under the influence of increased estrogen and progesterone, the uterine glands but become enlarged, contorted, and sacculated (having small lateral branches). Uterine glands function in the secretion of nutrient rich mucus fluid. • Blood vessels - spiral arteries – The blood vessels that deliver blood to the endometrium are the spiral arteries. The development of the spiral arteries is under hormonal control. Their growth is promoted by increased estrogen and progesterone levels. Atrophy of the spiral arteries occurs when levels of estrogen and progesterone become low. • Stroma – The stroma is the connective tissue that forms the framework of the endometrium, it contains the spiral arteries and the uterine glands. Figure 28.65 The endometrium is the inner mucosa of the uterus. It contains numerous uterine (endometrial) glands and blood vessels located within connective tissue called the stroma. The mucosa is located interior to the muscularis, the myometrium. 25 Endometrium • Stratum basalis – The stratum basalis is the layer of the endometrium close to the muscularis. The stratum basalis is the permanent layer of the endometrium. Uterine glands and blood vessels are maintained in the stratum basalis. • Stratum functionalis – The stratum functionalis is the layer of the endometrium closest to the uterine cavity and undergoes modifications in preparation of implantation of the fertilized egg. Uterine Cycle • The uterine (menstrual) cycle is divided into three phases, the (1) menstrual phase, (2) proliferative phase, and (3) secretory phase. • During the cycle the stratum functionalis, in response to changing levels of estrogen and progesterone, undergoes considerable structural changes. Uterine Cycle • Proliferative phase – The proliferative phase begins the rebuilding of the stratum functionalis. The proliferative phase begins in response to increasing levels of estrogen from the developing follicles (granulosa cells) of the ovary. Uterine Cycle The uterine (menstrual) cycle is divided into three phases, the (1) menstrual phase, (2) proliferative phase, and (3) secretory phase. Uterine Cycle • Menstrual Cycle – The menstrual cycle is the complete cycle of the uterus (of approximately 28 days) from one menstrual period to the next. • Menstrual phase (period) – The menstrual period is the time in the menstrual cycle in which menses, the flow of menstrual fluid, occurs. – The menstrual period occurs when levels of estrogen and progesterone are at their lowest, and is characterized by the shedding of the existing developed stratum functionalis. – This phase corresponds to the ending of the luteal phase of the ovarian cycle characterized by the degeneration of the corpus luteum with the reduction of progesterone (and estrogen) levels. Uterine Cycle • Secretory phase – The secretory phase continues the rebuilding with additional preparations such as the secretion of glycoproteins for receiving the fertilized egg. – The secretory phase is enhanced by increased levels of progesterone and estrogen. Progesterone and estrogen levels increase from the development of the corpus luteum following ovulation. – If implantation does not occur, the corpus luteum degenerates resulting in reduced levels of progesterone and estrogen. With reduced levels of hormones, the next uterine cycle begins with the menstrual phase. The menstrual phase results in the removal of the developed stratum functionalis. – If implantation occurs, the corpus luteum is maintained for a short time and the uterus continues to develop as the uterus of pregnancy and supports the developing embryo and fetus. 26 Uterine Cycle Figure 28.66 The uterine (menstrual) cycle is divided into three phases, the (1) menstrual phase, (2) proliferative phase, and (3) secretory phase. During the cycle the stratum functionalis, in response to changing levels of estrogen and progesterone, undergoes considerable structural changes. Lab Activity 14 – Menstrual Phase Figure 28.68 Scanning power photograph of the menstrual phase of the uterus. The endometrium is thin, and the stroma and uterine glands are filled with blood from ruptured blood vessels. Uterine Cycle Figure 28.67 Photographs of the phases of the uterus (human). The uterine cycle is divided into three phases, the (1) menstrual phase (menses), (2) proliferative phase, and (3) secretory phase. The phases may be further divided into early and late. A Menstrual Phase, Days 1 - 4 B Proliferative Phase, Days 4 - 14 C Early Secretory Phase, Days 14 - 18 D Secretory Phase, Days 18 - 23 E Secretory Phase, Days 23 - 25 F Late Secretory Phase, Days 25 - 28 G Pregnant uterus Lab Activity 15 – Proliferative Phase Figure 28.70 Scanning power photograph of the proliferative phase of the uterus (human), days 4-14. The uterine glands are observed as tubes with slight convolutions. Lab Activity 16 – Secretory Phase VAGINA The vagina is located between the cervix of the uterus and the opens at the external region between the labia minora, the vestibule Figure 28.71 Scanning power photograph of the late secretory phase of the uterus (human), days 23-25. The uterine glands are large and highly coiled with sacculations. 27 Vagina The vagina is located between the cervix of the uterus and the opens at the external region between the labia minora, the vestibule. • The vagina functions to receive the penis during intercourse and serves as the lower part of the birth canal. • The inner layer of the wall of the vagina is the mucosa and is lined with non-keratinized stratified squamous epithelium. The middle muscularis of the vagina is composed of circular and longitudinal layers of smooth muscle. Lab Activity 17 – Vagina • Figure 28.72 Scanning power photograph of the vagina (human). The inner layer of the vagina, the mucosa, is lined by non-keratinized stratified squamous epithelium. The muscularis consists of smooth muscle, an inner circular and an outer longitudinal layer. Lab Activity 17 – Vagina MAMMARY GLANDS Figure 28.73 Low power photograph of the stratified squamous epithelium of the mucosa. Mammary Glands • The paired mammary glands (commonly called breasts) are modified sebaceous glands that function in the production of milk, or lactation. • They are located within the superficial fascia on the anterior sides of the chest. Each mammary gland is associated with a protuberance called a nipple. • A nipple contains ducts from the glandular tissue that transport milk to the outside. Around each nipple is a pigmented area called the areola. The paired mammary glands (commonly called breasts) are modified sebaceous glands that function in the production of milk, or lactation. Mammary Glands • Mammary glands consist of glandular tissue surrounded and organized by fibrous connective tissue and fat. • The glandular tissue is organized into lobes, which are subdivided into lobules. The secretory units of the lobules are called alveoli, sacs of cells that are found at the terminal ends of excretory ducts. • The excretory ducts of the alveoli merge into small lacteriferous ducts. The lacteriferous ducts continue to merge and form a large lacteriferous duct for each lobe (glandular unit). The lacteriferous ducts converge to the base of the nipple and each expands to form a lacteriferous sinus. • The lacteriferous sinuses enter the nipple and open to the outside. 28 Mammary Glands Mammary Glands • During pregnancy the primary hormones for the development of the mammary glands include estrogen, progesterone, and prolactin. – Prolactin, secreted from the anterior pituitary is the primary hormone for the production of milk. – Milk is not produced prior to birth because estrogen and progesterone have an inhibitory effect on prolactin. However, following birth estrogen and progesterone levels decrease and their inhibitory effect on prolactin is removed. Figure 28.74 Mammary glands consists of glandular tissue surrounded and organized by fibrous connective tissue and fat. The glandular tissue is organized into lobes, which are subdivided into lobules. Each lobe is associated with a lactiferous duct, which expands near the nipple into a lacteriferous sinus. Mammary Glands Lab Activity 18 – Active Mammary Glands • Mammary glands that are hormonally stimulated during pregnancy undergo increased glandular development. • The lobules, undergo dramatic increases in the production and size of their secretory units, the alveoli. • Thus, mammary glands are described as active and inactive by the degree of glandular development and activity. Figure 28.75 Low power photograph of the well developed lobules of an active mammary gland. The secretory units of the lobules are called alveoli. Lab Activity 18 – Active Mammary Glands Figure 28.76 High power photograph of a lobule of an active mammary gland showing the secretory units called alveoli. Lab Activity 19 – Inactive Mammary Glands Figure 28.77 Low power photograph of the lobules of an inactive mammary gland. The connective tissue between the lobules, the interlobular septa, and associated fat appear abundant because of the reduced size of the alveoli. 29