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28-1 Structures of the Reproductive System
The Male Reproductive System Testes or male gonads Secrete male sex hormones (androgens) Produce male gametes (spermatozoa or sperm)
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Figure 28-4a The Structure of the Testes.
Ductus deferens Epididymis Mediastinum of testis Efferent ductule Rete testis Scrotum Skin Dartos muscle Straight tubule Superficial scrotal fascia Seminiferous tubules Cremaster muscle Tunica vaginalis Tunica albuginea Septa Scrotal cavity Septa Lobule Raphe A transverse section of the scrotum and testes a
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28-2 Male Reproductive Functions
Spermatogenesis Is the process of sperm production Begins at outermost cell layer in seminiferous tubules Proceeds toward lumen
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28-2 Male Reproductive Functions
Cells of Spermatogenesis Spermatogonia (stem cells) divide by mitosis to produce two daughter cells One remains as spermatogonium Second differentiates into primary spermatocyte Primary spermatocytes begin meiosis and form secondary spermatocytes
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28-2 Male Reproductive Functions
Cells of Spermatogenesis Secondary spermatocytes differentiate into spermatids (immature gametes) Spermatids Differentiate into spermatozoa Spermatozoa Lose contact with wall of seminiferous tubule Enter fluid in lumen
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Figure 28-5c The Seminiferous Tubules.
Spermatid Nurse cell Dividing spermatocytes Capillary Lumen Interstitial cells Spermatogonium Spermatozoa c c Nurse cells surround the stem cells of the tubule and support the developing spermatocytes and spermatids.
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Figure 28-5d The Seminiferous Tubules.
Spermatids completing spermiogenesis LUMEN Spermatids beginning spermiogenesis Initial spermiogenesis Secondary spermatocyte Secondary spermatocyte in meiosis II Primary spermatocyte preparing for meiosis I Luminal compartment Blood–testis barrier Nuclei of nurse cells Fibroblast Connective tissue capsule Capillary Interstitial cells Spermatogonium Basal compartment d Stages in spermatogenesis in the wall of a seminiferous tubule. d
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28-2 Male Reproductive Functions
Spermatogenesis Involves three integrated processes Mitosis Meiosis Spermiogenesis
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28-2 Male Reproductive Functions
Mitosis Is part of somatic cell division Produces two diploid (2n) daughter cells Both have identical pairs of chromosomes
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Figure 28-6a Chromosomes in Mitosis and Meiosis.
Mitosis. The fates of three representative chromosome pairs during mitosis Cell (2n) Chromosomes Chromosome duplication 1 2 Chromosomes of daughter cell 1 Chromosomes of daughter cell 2 Daughter cells (2n)
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Figure 28-6b Chromosomes in Mitosis and Meiosis.
Meiosis. The fates of three representative chromosome pairs during the two stages of meiosis Cell (2n) Chromosomes Tetrad Chromosome duplication, synapsis, and tetrad formation Meiosis I Meiosis II 1 2 3 4 1 2 3 4 Chromosomes of gametes Gametes (n)
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Figure 28-7 Spermatogenesis (Part 4 of 4).
Spermiogenesis (physical maturation) In spermiogenesis, the last step of spermatogenesis, each spermatid matures into a single spermatozoon, or sperm. The process of spermiogenesis—from spermatids to spermatozoa—takes 24 days. Spermatozoa (haploid, n)
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28-2 Male Reproductive Functions
Six Major Functions of Nurse Cells Maintain blood–testis barrier Support mitosis and meiosis Support spermiogenesis Secrete inhibin Secrete androgen-binding protein (ABP) Secrete Müllerian-inhibiting factor (MIF)
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28-2 Male Reproductive Functions
Secretion of Inhibin Inhibin Is a peptide hormone secreted by nurse cells in response to factors released by spermatozoa Depresses: Pituitary production of FSH Hypothalamic secretion of GnRH Regulation of FSH and GnRH by inhibin: Gives nurse cells feedback control of spermatogenesis After division, increases inhibin production
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28-2 Male Reproductive Functions
Secretion of Androgen-Binding Protein (ABP) Androgen-Binding Protein (ABP) Binds androgens (primarily testosterone) In seminiferous tubule fluid Is important in: Elevating androgen in seminiferous tubules Stimulating spermiogenesis Production of ABP is stimulated by FSH
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28-2 Male Reproductive Functions
Secretion of Müllerian-Inhibiting Factor (MIF) Müllerian-Inhibiting Factor (MIF) Is secreted by nurse cells in developing testes Causes regression of fetal Müllerian (paramesonephric) ducts Help form uterine tubes and uterus in females In males, inadequate MIF production leads to: Retention of ducts Failure of testes to descend into scrotum
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Figure 28-8 Spermiogenesis and Spermatozoon Structure (Part 2 of 2).
Tail (55 m) Fibrous sheath of flagellum Middle piece (5 m) Spermatozoa SEM × 780 Mitochondrial spiral Neck (1 m) Centrioles Nucleus Head (5 m) Acrosome Spermatozoon
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28-2 Male Reproductive Functions
The Male Reproductive Tract Sperm maturation Testes produce physically mature spermatozoa that CANNOT fertilize an oocyte Other parts of reproductive system are responsible for: Functional maturation, nourishment, storage, and transport
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28-2 Male Reproductive Functions
The Male Reproductive Tract Sperm maturation Spermatozoa Detach from nurse cells Are free in lumen of seminiferous tubule Are functionally immature Are incapable of locomotion or fertilization Are moved by cilia lining efferent ductules into the epididymis
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Figure 28-9a The Epididymis.
Spermatic cord Ductus deferens Efferent ductules Straight tubule Rete testis Seminiferous tubule Epididymis Head of epididymis Tunica albuginea Body of epididymis Testis Tail of epididymis Scrotal cavity a A diagrammatic view of the head, body, and tail of an epididymis
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28-2 Male Reproductive Functions
Three Functions of the Epididymis Monitors and adjusts fluid produced by seminiferous tubules Recycles damaged spermatozoa Stores and protects spermatozoa Facilitates functional maturation
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28-2 Male Reproductive Functions
Two Steps in Capacitation Spermatozoa become motile When mixed with secretions of seminal glands Spermatozoa become capable of fertilization When exposed to female reproductive tract
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Figure 28-10a The Ductus Deferens and Accessory Glands.
Ureter Urinary bladder Ductus deferens Seminal gland Ampulla of ductus deferens Duct of seminal gland Ejaculatory duct Prostate gland Prostatic urethra a A posterior view of the urinary bladder and prostate gland, showing subdivisions of the ductus deferens in relation to surrounding structures. Bulbourethral glands Urogenital diaphragm
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28-2 Male Reproductive Functions
Four Major Functions of Male Glands Activating spermatozoa Providing nutrients spermatozoa need for motility Propelling spermatozoa and fluids along reproductive tract Mainly by peristaltic contractions Producing buffers To counteract acidity of urethral and vaginal environments
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28-2 Male Reproductive Functions
Semen Typical ejaculation releases 2–5 mL Abnormally low volume may indicate problems With prostate gland or seminal glands Sperm count Is taken of semen collected after 36 hours of sexual abstinence Normal range 20–100 million spermatozoa/mL of ejaculate
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28-2 Male Reproductive Functions
Ejaculate Is the volume of fluid produced by ejaculation Contains: Spermatozoa Seminal fluid Enzymes Including protease, seminalplasmin, prostatic enzyme, and fibrinolysin
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Figure 28-12 Regulation of Male Reproduction (Part 1 of 2).
Male reproductive function is regulated by the complex interaction of hormones from the hypothalamus, anterior lobe of the pituitary gland, and the testes. Negative feedback systems keep testosterone levels within a relatively narrow range until late in life. HYPOTHALAMUS Release of Gonadotropin- Releasing Hormone (GnRH) Negative feedback ? High testosterone levels inhibit the release of GnRH by the hypothalamus, causing a decrease in LH secretion, which lowers testosterone to normal levels. The hypothalamus secretes the hormone GnRH at a rate that remains relatively steady. As a result, blood levels of FSH, LH, and testosterone remain within a relatively narrow range throughout a man’s reproductive life. When stimulated by GnRH, the anterior lobe of the pituitary gland releases luteinizing hormone (LH) and follicle-stimulating hormone (FSH). ANTERIOR LOBE OF THE PITUITARY GLAND Secretion of Follicle- Stimulating Hormone (FSH) Secretion of Luteinizing Hormone (LH) Inhibin depresses the pituitary production of FSH, and perhaps the hypothalamic secretion of gonadotropin-releasing hormone (GnRH).The faster the rate of sperm production, the more inhibin is secreted. By regulating FSH and GnRH secretion, nurse cells provide feedback control of spermatogenesis. FSH targets primarily the nurse cells of the seminiferous tubules. LH targets the interstitial cells of the testes. KEY Stimulation Inhibition
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Figure 28-12 Regulation of Male Reproduction (Part 2 of 2).
Interstitial Cell Stimulation Negative feedback LH induces the secretion of testosterone and other androgens by the interstitial cells of the testes. TESTES Nurse Cell Stimulation Testosterone Under FSH stimulation, and with testosterone from the interstitial cells, nurse cells (1) secrete inhibin in response to factors released by developing spermatozoa, (2) secrete androgen-binding protein (ABP), and (3) promote spermatogenesis and spermiogenesis. Inhibin KEY Stimulation Inhibition Androgen-binding protein (ABP) binds androgens within the seminiferous tubules, which increases the local concentration of androgens and stimulates the physical maturation of spermatids. Nurse cell environment facilitates both spermatogenesis and spermiogenesis. Peripheral Effects of Testosterone Maintains libido (sexual drive) and related behaviors Stimulates bone and muscle growth Establishes and maintains male secondary sex characteristics Maintains accessory glands and organs of the male reproductive system
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Figure 28-14b The Ovaries and Their Relationships to the Uterine Tube and Uterus.
Corpus luteum Cortex Tunica albuginea Mesenteries of the Ovary and Uterine Tube Mesosalpinx Mesovarium Broad ligament Ovarian hilum Egg nest Germinal epithelium Mature follicle b A sectional view of the ovary, uterine tube, and associated mesenteries
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28-3 The Female Reproductive System
Oogenesis Also called ovum production Begins before birth Accelerates at puberty Ends at menopause
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Figure 28-15 Oogenesis (Part 1 of 3).
Mitosis of oogonium Oogonium (stem cell) Unlike spermatogonia, the oogonia (ō-ō-GŌ-nē-uh), or female reproductive stem cells, complete their mitotic divisions before birth. Oogonium Primary oocyte (diploid, 2n)
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Figure 28-15 Oogenesis (Part 2 of 3).
Meiosis I DNA replication Between the third and seventh months of fetal development, the daughter cells, or primary oocytes (Ō-ō-sīts), prepare to undergo meiosis. They proceed as far as the prophase of meiosis I, but then the process comes to a halt. Primary oocyte Tetrad First polar body The primary oocytes remain in a state of suspended development until puberty, when rising levels of FSH trigger the start of the ovarian cycle. Each month after the ovarian cycle begins, some of the primary oocytes are stimulated to undergo further development. Meiosis I is then completed, yielding a secondary oocyte and a polar body. Secondary oocyte First polar body may not complete meiosis II
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Figure 28-15 Oogenesis (Part 3 of 3).
Meiosis II Secondary oocyte released (ovulation) in metaphase of meiosis II Each month after the ovarian cycle begins, one or more secondary oocytes leave the ovary suspended in metaphase of meiosis II. At the time of fertilization, a second polar body forms and the fertilized oocyte is then called a mature ovum. (A cell in any of the preceding steps in oogenesis is sometimes called an immature ovum.) Second polar body Sperm (n) Nucleus of oocyte (n) Fertilization (see Figure 29–1)
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28-3 The Female Reproductive System
Fetal Development Between third and seventh months: Primary oocytes prepare for meiosis Stop at prophase of meiosis I Atresia Is the degeneration of primordial follicles Ovaries have about 2 million primordial follicles at birth Each containing a primary oocyte By puberty: Number drops to about 400,000
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28-3 The Female Reproductive System
Oogenesis: Two Characteristics of Meiosis Cytoplasm of primary oocyte divides unevenly Producing one ovum (with original cytoplasm) And two or three polar bodies (that disintegrate) Ovary releases secondary oocyte (not mature ovum) Suspended in metaphase of meiosis II Meiosis is completed upon fertilization
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28-3 The Female Reproductive System
The Ovarian Cycle After sexual maturation A different group of primordial follicles is activated each month Is divided into: Follicular phase (preovulatory phase) Luteal phase (postovulatory phase)
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28-3 The Female Reproductive System
The Ovarian Cycle Ovarian follicles Are specialized structures in cortex of ovaries Where oocyte growth and meiosis I occur Primary oocytes Are located in outer part of ovarian cortex Near tunica albuginea In clusters called egg nests
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28-3 The Female Reproductive System
Primordial Follicle Each primary oocyte in an egg nest Is surrounded by follicle cells Primary oocyte and follicle cells form a primordial follicle
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Figure 28-16 The Ovarian Cycle (Part 1 of 7).
Primordial follicles in egg nest Primary oocyte Follicle cells LM ×1440
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Figure 28-16 The Ovarian Cycle (Part 2 of 7).
Formation of primary follicle Granulosa cells Primary oocytes Zona pellucida Thecal cells LM ×1092
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Figure 28-16 The Ovarian Cycle (Part 3 of 7).
Formation of secondary follicle Thecal cells Zona pellucida Nucleus of primary oocyte Granulosa cells LM ×1052
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Figure 28-16 The Ovarian Cycle (Part 4 of 7).
Formation of tertiary follicle Antrum containing follicular fluid Granulosa cells Corona radiata Secondary oocyte LM ×136
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Figure 28-16 The Ovarian Cycle (Part 5 of 7).
Primordial follicles Primary follicle Secondary follicle Tertiary follicle Released secondary oocyte Corona radiata Corpus albicans Corpus luteum 5 Ovulation Follicular fluid Secondary oocyte within corona radiata Ruptured follicle wall Outer surface of ovary LM ×70
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28-3 The Female Reproductive System
The Uterine Cycle (Menstrual Cycle) Is a repeating series of changes in endometrium Lasts from 21 to 35 days Average 28 days Responds to hormones of ovarian cycle Menses and proliferative phase Occur during ovarian follicular phase Secretory phase Occurs during ovarian luteal phase
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28-3 The Female Reproductive System
Menses Is the degeneration of functional zone Occurs in patches Is caused by constriction of spiral arteries Reducing blood flow, oxygen, and nutrients Weakened arterial walls rupture Releasing blood into connective tissues of functional zone
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28-3 The Female Reproductive System
Menses Degenerating tissues break away, enter uterine lumen Entire functional zone is lost Through external os and vagina Only functional zone is affected Deeper, basilar zone is supplied by straight arteries
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The endometrium at menses
Figure 28-20a The Appearance of the Endometrium during the Uterine Cycle. Perimetrium Uterine glands Uterine cavity Endometrium Myometrium Basilar zone of endometrium Cervix MYOMETRIUM Menses LM ×63 a The endometrium at menses
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28-3 The Female Reproductive System
The Proliferative Phase Epithelial cells of uterine glands Multiply and spread across endometrial surface Restore integrity of uterine epithelium Further growth and vascularization Completely restores functional zone Occurs at same time as: Enlargement of primary and secondary follicles in ovary
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28-3 The Female Reproductive System
The Proliferative Phase Is stimulated and sustained by: Estrogens secreted by developing ovarian follicles Entire functional zone is highly vascularized Small arteries Spiral toward inner surface From larger arteries in myometrium
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The endometrium during the proliferative phase
Figure 28-20b The Appearance of the Endometrium during the Uterine Cycle. Uterine glands Uterine cavity Uterine cavity Functional zone Myometrium ENDOMETRIUM Basilar zone MYOMETRIUM Proliferative phase LM ×66 b The endometrium during the proliferative phase
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28-3 The Female Reproductive System
The Secretory Phase Endometrial glands enlarge, increasing rate of secretion Arteries of uterine wall Elongate and spiral through functional zone Begins at ovulation and persists as long as corpus luteum remains intact Peaks about 12 days after ovulation Glandular activity declines Generally lasts 14 days Ends as corpus luteum stops producing stimulatory hormones
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The endometrium during the secretory phase of the uterine cycle
Figure 28-20c The Appearance of the Endometrium during the Uterine Cycle. Uterine glands Uterine cavity Detail of uterine glands LM ×150 Functional zone Secretory phase LM ×52 c The endometrium during the secretory phase of the uterine cycle
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Figure 28-24 Regulation of Female Reproduction (Part 3 of 6).
The ovarian and uterine cycles must operate in synchrony to ensure proper reproductive function. If the two cycles are not properly coordinated, infertility results. A female who doesn’t ovulate cannot conceive, even if her uterus is perfectly normal. A female who ovulates normally, but whose uterus is not ready to support an embryo, will also be infertile. HYPOTHALAMUS As in males, GnRH from the hypothalamus regulates reproductive function in females. However, in females, GnRH levels change throughout the course of the ovarian cycle. 1 Release of Gonadotropin-Releasing Hormone (GnRH) The cycle begins with the release of GnRH, which stimulates the production and secretion of FSH and the production—but not the secretion— of LH. KEY Stimulation Release of GnRH Inhibition ANTERIOR LOBE OF PITUITARY GLAND 2 3 Follicular Phase of the Ovarian Cycle Luteal Phase of the Ovarian Cycle Production and secretion of FSH Production of LH The follicular phase begins when FSH stimulates some secondary follicles to develop into a tertiary follicle. The combination of increased GnRH pulse frequency and elevated estrogen levels stimulates LH secretion. Secretion of LH On or around day 14, a massive surge in LH level triggers (1) the completion of meiosis I by the primary oocyte, (2) the forceful rupture of the follicular wall, (3) ovulation, roughly 9 hours after the LH peak, and (4) formation of the corpus luteum. As secondary follicles develop, FSH levels decline due to the negative feedback effects of inhibin. Negative feedback
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Figure 28-24 Regulation of Female Reproduction (Part 4 of 6).
Follicular Phase of the Ovarian Cycle Developing follicles also secrete estrogens, especially estradiol, the dominant hormone prior to ovulation. 3 Luteal Phase of the Ovarian Cycle OVARY After day 10 The corpus luteum secretes progesterone, which stimulates and sustains endometrial development. Before day 10 • Meisois I completion • Ovulation • Corpus luteum formation In low concentrations, estrogens inhibit LH secretion. This inhibition gradually decreases as estrogen levels increase. • Follicle development • Secretion of inhibin estrogens After ovulation, progesterone levels rise and estrogen levels fall. This suppresses GnRH secretion. If pregnancy does not occur, the corpus luteum will degenerate after 12 days, and as progesterone levels decrease, GnRH secretion increases, and a new cycle begins. Secretion of progesterone Effects on CNS Stimulation of bone and muscle growth Establishment and maintenance of female secondary sex characteristics Maintenance of accessory glands and organs Stimulation of endometrial growth and secretion KEY Stimulation Inhibition
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Figure 28-24 Regulation of Female Reproduction (Part 5 of 6).
FOLLICULAR PHASE OF OVARIAN CYCLE LUTEAL PHASE OF OVARIAN CYCLE 50 40 LH Gonadotropic hormone levels (IU/L) 30 GnRH pulse frequency (pulses/day) 20 FSH 10 Follicle stages during the ovarian cycle Corpus luteum formation Mature corpus luteum Corpus albicans Follicle development Ovulation Progesterone Ovarian hormone levels Estrogens Inhibin 28/0 7 14 21 28/0 Days
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Figure 28-24 Regulation of Female Reproduction (Part 6 of 6).
FOLLICULAR PHASE OF OVARIAN CYCLE LUTEAL PHASE OF OVARIAN CYCLE Destruction of functional zone Repair and regeneration of functional zone Secretion by uterine glands Endometrial change during thes uterine cycle Phases of the uterine cycle MENSES PROLIFERATIVE PHASE SECRETORY PHASE 36.7 Basal body temperature (C) 36.4 28/0 7 14 21 28/0 Days
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28-3 The Female Reproductive System
Summary: Hormonal Regulation of the Female Reproductive Cycle Early in follicular phase of ovarian cycle: Estrogen levels are low GnRH pulse frequency is 16–24/day (1 per 60–90 minutes) As tertiary follicles form, concentration of circulating estrogens rises steeply And GnRH pulse frequency increases to 36/day (1 per 30–60 minutes)
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28-3 The Female Reproductive System
Summary: Hormonal Regulation of the Female Reproductive Cycle In follicular phase: Switchover occurs When estrogen levels exceed threshold value for about 36 hours Resulting in massive release of LH from the anterior lobe of the pituitary gland
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28-3 The Female Reproductive System
Summary: Hormonal Regulation of the Female Reproductive Cycle In follicular phase: Sudden surge in LH concentration triggers: Completion of meiosis I by primary oocyte Rupture of follicular wall Ovulation Ovulation occurs 34–38 hours after LH surge begins (nine hours after LH peak)
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28-3 The Female Reproductive System
Summary: Hormonal Regulation of the Female Reproductive Cycle In luteal phase of ovarian cycle: High LH levels trigger ovulation Promote progesterone secretion Trigger formation of corpus luteum Frequency of GnRH pulses stimulates LH more than FSH LH maintains structure and secretory function of corpus luteum
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28-3 The Female Reproductive System
Summary: Hormonal Regulation of the Female Reproductive Cycle Luteal phase Progesterone levels remain high for one week Unless pregnancy occurs, corpus luteum begins to degenerate Progesterone and estrogen levels drop GnRH pulse frequency increases Stimulating FSH secretion Ovarian cycle begins again
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28-3 The Female Reproductive System
Hormones and the Uterine Cycle Corpus luteum degenerates Progesterone and estrogen levels decline Resulting in menses Endometrial tissue sheds several days Until rising estrogen stimulates regeneration of functional zone
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28-3 The Female Reproductive System
Hormones and the Uterine Cycle Proliferative phase continues Until rising progesterone starts secretory phase Increase in estrogen and progesterone Causes enlargement of endometrial glands And increase in secretory activities
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28-3 The Female Reproductive System
Hormones and Body Temperature Monthly hormonal fluctuations affect core body temperature During luteal phase, progesterone dominates During follicular phase, estrogen dominates and basal body temperature decreases about 0.3C Upon ovulation, basal body temperature (BBT) declines noticeably Day after ovulation, temperature rises
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