Gonads
Gonads refer to the testes and ovaries present in male and female animals, respectively. They are responsible for the production of gametes and hormones, which are crucial elements governing sexual reproduction in animals (Table 17.4).
The hypothalamic-pituitary axis secretes GnRH, LH, and FSH, which are essential for regulating the endocrine and gametogenic activities of gonads. In turn, the gonadal hormones produced exert negative inhibition on the release of hypothalamic GnRH and pituitary gonadotropins.17.6.1 Ovary
The follicle and corpus luteum developed sequentially are the transient endocrine organs that secrete two major steroids: estrogen and progesterone, respectively. The granulosa cells present in the follicle produce estrogen from androgens synthesised in the theca cells. Estrogen plays a vital role in the estrous cycle, development of female secondary sexual characteristics, ductular growth in the mammary gland, and parturition. Apart from the steroidal hormones, peptide hormones such as inhibin, activin, follistatin, and anti-Mullerian hormone are produced from the ovarian follicle. Whereas progesterone secretion to support gestation and lobuloalveolar development in the mammary gland is the major endocrine function of the corpus luteum. In addition, it also secretes relaxin and oxytocin that are responsible for the widening of the birth canal and luteal demise in cyclical animals.
Table 17.4 List of gonadal hormones and their functions
| S. No | Hormone | Chemical nature | Function |
| 1. | Estrogen (Females: Granulosa cell, Males: Sertoli cell) | Steroid (18C) | • Estrus cycle • Female sexual behavior • Parturition • Ductular growth in mammary gland |
| 2. | Inhibin (Females: Granulosa cell, Males: Sertoli cell) | Glycoprotein | • Inhibit the release of FSH from the anterior pituitary gland |
| 3. | Activin (Follicular fluid and Rete testis fluid) | Glycoprotein | • Stimulates the secretion of FSH from the anterior pituitary gland |
| 4. | Follistatin | Protein | • Inhibits FSH secretion • Neutralizes activin |
| 5. | Testosterone (Females: Theca cell, Males: Leydig cell) | Steroid (19C) | • Masculine characters in male animals • Libido • Protein anabolism • Development of accessory sex glands |
| 6. | Progesterone (Corpus luteum) | Steroid (21C) | • Maintenance of pregnancy • Lobuloalveolar growth of the mammary gland |
| 7. | Anti-Mullerian hormone | Protein | • Regression of Mullerian duct |
17.6.2 Testes
The Sertoli and Leydig cells are the somatic cell types with endocrine function in testes.
The testosterone produced from the Leydig cells in response to LH imparts secondary sexual characteristics, anabolic effects on bone and muscle mass, and libido in male animals. Additionally, the Sertoli cells are responsible for the secretion of inhibin, which attenuates the secretion of FSH from the adenohypophysis.Endocrinology to Rescue Wild Animals...
• Several wild animal species are staring at an impending threat of extinction more than ever in the history. The ex situ breeding and managemental practices play a huge role in reviving their population and curtail them from getting extinct. Endocrinological approaches such as measuring the plasma levels of hormones such as estrogen, progesterone, and cortisol help in assessing the reproductive status and well-being of different wild animals. They also greatly contribute in understanding the reproductive behavior of animals and aid in devising intervening strategies of breeding in captive wild animals.
Learning Outcomes
• Natriuretic peptides: ANP, BNP, and CNP are the major natriuretic peptides secreted from cardiac tissue and CNS. They are secreted due to the stretch of cardiac cells due to an increased blood volume. The natriuretic peptides (ANP and BNP) are responsible for the excretion of sodium and water from kidneys to regulate the circulatory volume to a normal level. They cause an increase in the glomerular filtration via their vasodilatory effect on afferent arteriole. Decreased tubular absorption of sodium and water leads to an increased excretion with a simultaneous decrease in the blood volume. The natriuretic and diuretic effects of natriuretic peptides antagonize the antidiuretic action of aldosterone, which is known as the aldosterone escape.
• Erythropoietin: It is a glycoprotein hormone released from the kidneys in response to tissue hypoxia. The bone marrow as its target organ, EPO acts on the erythropoietic stem cells to increase their proliferation, differentiation, and subsequent transformation into the erythrocytes.
• Thymus: A major organ determines the cell- mediated immunity in animals by acting as the site of differentiation and production of T-cells. Additionally, the thymic epithelial cells possess the endocrine activity and secrete hormones such as thymulin, thymopoietin, and thymosins. These hormones primarily act to regulate the activation and differentiation of T-cells and modulate the immune status of the animals.
• Gut hormones: The enteroendocrine cells present in the GIT produce a wide variety of peptide hormones. The enteroendocrine cells act as nutrient sensors and secrete the hormones that control the appetite, gastrointestinal motility, exocrine secretion of the pancreas, liver, and GIT. All these hormones work in concert to harvest appropriate amounts of nutrients by tightly regulating the process of digestion and metabolism.
• Placenta: Recognized as a crucial transient organ formed during the early gestation period and responsible for supplying the nutrients to the fetus. Furthermore, it also secretes important hormones such as progesterone, estrogen, chorionic gonadotropins, placental lactogen, and relaxin. Together, they ensure the maintenance of pregnancy, fetal growth, and progression of parturition.
• Gonads: The production of gametes and hormones that control sex-specific behaviors in animals are two important functions of gonads. The testes in male animals produce testosterone, responsible for libido, secondary sexual characteristic, anabolic effects on bone and muscles. Estrogen is the major circulatory steroid produced by the ovarian follicles responsible for reproductive cyclicity, increasing bone mass, growth of reproductive tract and mammary gland in female animals. In addition, corpus luteum formed after ovulation produces progesterone, which helps in the growth of endometrium and immunotolerance to support the fetal growth in the uterus.
Exercises
Objective Questions
Q1. ___________ is a natriuretic peptide that cannot pro
mote the excretion of sodium
Q2.
Natriuretic peptides activate______________ enzymeto initiate the signal transduction.
Q3. The antagonistic effects of natriuretic peptides on aldosterone are collectively known as
Q4. The secondary messenger produced in the target cells when bound with natriuretic peptides is
Q5. Erythropoietin is a____________ type of hormone, and
released primarily due to_______________.
Q6. The thymic hormones are produced by cells present in the thymus
Q7. Which thymic hormone requires zinc for its biological activity?
Q8. What is the first hormone to be discovered?
Q9. The cells in GIT that produce gut hormones are collectively called as
Q10. Which hormone is responsible for migrating myoelectric complex?
Q11. The gut hormones that stimulate insulin secretion from the pancreas are known as
Q12. Which hormone is responsible for the relaxation of pelvic symphysis before parturition?
Q13. The specialized areas in equine placenta that secrete eCG are called as
Q14. Which placental hormone has structural homology of both GH and PRL?
Q15. Which gonadal hormone is responsible for the lobuloalveolar development of the mammary gland?
Subjective Questions
Q1. Enlist the different types of natriuretic peptides and their major production sites.
Q2. Briefly describe the biological effects of ANP.
Q3. What is EPO? Describe its effect on the bone marrow.
Q4. Explain the mechanism of production of EPO from kidneys.
Q5. Describe briefly the endocrine role of the thymus
Q6. What are the orexigenic hormones produced by the GIT?
Q7. What are the biological effects of gut hormones?
Q8. Enlist various placental hormones and briefly describe their biological effects in animals
Q9. What are the cyclical endocrine structures developed in the ovary and the respective hormones secreted from them?
Q10. What are the different somatic cells present in the testes and list their hormones?
Answers to Objective Questions
A1.
CNPA2. Guanylyl cyclase
A3. Aldosterone escape
A4. cGMP
A5. Glycoprotein, tissue hypoxia
A6. Thymic epithelial cells
A7. Thymulin
A8. Secretin
A9. Enteroendocrine cells
A10. Motilin
A11. Incretins
A12. Relaxin
A13. Endometrial cups
A14. Placental lactogen
A15. Progesterone
Answers to Subjective Questions
A1. ANP, BNP, CNP, cardiac muscle, and CNS
A2. Natriuresis, diuresis, vasodilation in arterioles, and inhibit aldosterone secretion
A3. Erythropoietin, glycoprotein, kidney, and erythropoiesis
A4. Tissue hypoxia, HIF1α, oxygen-sensitive
hydroxylases, and hypoxia-response element
A5. Thymic epithelial cells, thymulin, thymopoietin, and thymosins
A6. Ghrelin, motilin, and insulin-like peptide 5 (INSL5)
A7. Regulating GIT motility, exocrine pancreatic secretion, gastric acid secretion, and appetite
A8. Progesterone, estrogen, placental lactogen, chorionic gonadotropins, and relaxin
A9. Follicle, corpus luteum, estrogen, and progesterone
A10. Sertoli cells-estrogen, inhibin; Leydig cells- testosterone
Further Reading
Textbooks
Hafez ESE, Hafez B (eds) (2013) Reproduction in farm animals. Wiley McDonald LE, Pineda MH, Dooley MP (2003) McDonald’s veterinary endocrinology and reproduction. 5th ed./edited by M.H. Pineda, with the editorial assistance of Michael P. Dooley. Ames, Iowa: Iowa State Press. Print
Norman AW, Henry HL (2022) Hormones. Academic Press. https://doi. org/10.1016/B978-0-08-091906-5.00007-0
Takei Y, Ando H, Tsutsui K (eds) (2015) Handbook of hormones: comparative endocrinology for basic and clinical research. Academic Press. https://doi.org/10.1016/C2013-0-15395-0
Articles
Ahlman H, Nilsson O (2001) The gut as the largest endocrine organ in the body. Ann Oncol 12:S63-S68. https://doi.org/10.1093/annonc/ 12.suppl_2.S63
Alvarez-Oxiley AV, De Sousa NM, Beckers JF (2008) Native and recombinant bovine placental lactogens. Reprod Biol 8(2):85-106. https://doi.org/10.1016/S1642-431X(12)60006-0
Goetze JP, Bruneau BG, Ramos HR, Ogawa T, de Bold MK, de Bold AJ (2020) Cardiac natriuretic peptides. Nat Rev. Cardiol 17(11):698-717. https://doi.org/10.1038/s41569-020-0381-0
Hall CM, Glaser S, Alpini G (2017) Gastrointestinal Hormone (GI) Regulated Signal Transduction. https://doi.org/10.1016/B978- 0-12-809324-5.03384-8
Latorre R, Sternini C, De Giorgio R, Greenwood-Van Meerveld B (2016) Enteroendocrine cells: a review of their role in brain-gut communication. Neurogastroenterol Motil 28(5):620-630. https:// doi.org/10.1111/nmo.12754
Savino W, Dardenne M (2000) Neuroendocrine control of thymus physiology. Endocr Rev 21(4):412-443. https://doi.org/10.1210/ edrv.21.4.0402
Schuler G, Furbass R, Klisch K (2018) Placental contribution to the endocrinology of gestation and parturition. Anim Reprod 15(Suppl 1):822-842. https://doi.org/10.5167/uzh-167451