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The Hypothalamic Releasing Hormones
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Thyrotropin-Releasing Hormone (TRH) This was the rst of the releasing hormones to be identi ed; its tripeptide structure was determined in 1968 The hormone is elaborated by the anterior periventricular, paraventricular, arcuate, ventromedial, and dorsomedial neurons but not by those of the posterior hypothalamic or thalamic nuclei Mainly, it stimulates the release of thyroidstimulating hormone (TSH) from the pituitary gland; TSH, in turn, effects the release of T4 (thyroxine) and T3 (triiodothyronine) Pituitary cells that release dopamine and somatostatin are also stimulated to a slight degree; the latter has an inhibitory effect on TSH There is also an inhibitory feedback of T3 on TSH and TRH Actually, more than half of brain TRH is found outside the hypothalamus in brainstem raphe nuclei, tractus solitarius, and the anterior and lateral horn cells of spinal cord suggesting that TRH may function as a central regulator of the autonomic nervous system Growth Hormone Releasing Hormone (GHRH) This hormone and somatostatin (also known as GH-release inhibiting hormone, or SRIH) are both secreted by specialized tuberoinfundibular neurons and released into the hypophysial-portal circulation, by which they are carried to speci c GH-secreting cells of the anterior pituitary gland (somatotropes) Immunohistochemical staining has shown the sources of GHRH and somatostatin to be neurons of the posterior part of the arcuate and ventromedian hypothalamic nuclei and other neurons of the median eminence and premammillary area Somatostatin, a 14 amino acid peptide, is produced more anteriorly by neurons in the periventricular area and small-cell part of the paraventricular nucleus The amygdala, hippocampus, and other limbic structures project to the arcuate nuclei via the medial
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peptides thyroid-releasing factor, vasoactive intestinal peptide, peptide-histidine-isoleucine, and oxytocin also have the capacity to raise the levels of prolactin in the blood The hypothalamopituitary axis is responsive to sensory stimuli from the nipples, via pathways in the spinal cord and brainstem, accounting for the effect of suckling on milk production The normal blood levels of prolactin are 5 to 25 ng/mL The release of prolactin is also in uenced by an inhibitory dopaminergic innervation of pituitary lactotrophic cells This latter mechanism accounts for galactorrhea and reproductive disorders, which occur with tumors that compress the pituitary stalk and interrupt the venous portal transport of dopamine from the hypothalamus
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The Neurohypophysis: Vasopressin and Oxytocin
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The oliogopeptides vasopressin and oxytocin are elaborated by cells of the supraoptic and paraventricular nuclei and are transported, via their axons, through the stalk of the pituitary to its posterior lobe, where these substances are stored Together, these elements constitute the neurohypophysis (posterior pituitary), which develops as an evagination of the oor of the third ventricle Some of the vasopressin-containing nerve endings also terminate on cells of origin of the autonomic nervous system and on the capillary plexus of the hypophysial portal circulation, through which they in uence the secretion of CRH and GH The peptide parts of vasopressin and oxytocin, whose chemical nature was determined by duVigneaud, are almost identical, differing from one another by only two amino acids Vasopressin, acting on the kidney tubules, serves as the antidiuretic hormone (ADH) and, complemented by thirst mechanisms, maintains the osmolality of the blood Plasma osmolality modi es vasopressin secretion by acting directly on the supraoptic and paraventricular neurons or on separate osmoreceptors in the hypothalamus The sensitivity of the vasopressin-ADH mechanism is demonstrated by the absence of antidiuretic effect when vasopressin is below 1 pg/mL and by maximal antidiuresis when plasma levels reach 5 pg/mL If serum osmolality falls below 280 mosm/L, the release of ADH is completely inhibited This system is most effective in maintaining homeostasis when serum osmolality is relatively close to the normal range, between 280 and 295 mosm/L Alterations in blood volume and pressure also affect vasopressin release through neural mechanisms that have their origin in baro- and mechanoreceptors of the aortic arch, carotid sinus, and right atrium Afferent signals from these regions are conveyed in the vagus and glossopharyngeal nerves, which synapse in the nucleus of the tractus solitarius; the precise pathways to the hypothalamus have not been delineated, however With severe hypotension, ADH release will continue even if there is a low serum osmolality, ie, pressure predominates over osmolarity as a stimulus Vasopressin secretion is also in uenced by nonosmotic factors Nausea, for example, is a potent stimulus, raising levels of the hormone 100-fold Hypoglycemia has a less profound effect Drugs such as morphine, nicotine, alcohol, and certain chemotherapeutic agents (cyclophosphamide) also cause release of the stored peptide Pain, emotional stress, and exercise have long been thought to cause release of vasopressin, but it is unclear whether this is a direct effect or is mediated through hypotension or nausea Oxytocin initiates uterine contraction and has milk-ejecting effects Its release is stimulated by distention of the cervix, labor,
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breast-feeding, and estrogen The effects of oxytocin are inhibited by alcohol In summary, it is apparent that the regulatory system of hypothalamic releasing hormones is complex The releasing factors have overlapping functions, and the hypothalamic nuclei act on many parts of the brain in addition to the pituitary Conversely, many parts of the brain in uence the hypothalamus through neural connections or modulate its activity and that of the pituitary gland through the action of neurotransmitters and modulators (catecholamines, acetylcholine, serotonin, and dopamine) There is feedback control between every part of the hypothalamus and the endocrine structures on which it acts The factors that in uence hypothalamic neurons have been reviewed in detail by Reichlin Some of these relationships have been mentioned and others will emerge further on in this chapter and in later chapters, particularly as they relate to behavioral and psychiatric disorders Of particular signi cance is the role of the hypothalamus in the integration of the endocrine and autonomic nervous systems at both the peripheral and central levels The best-known example of this interaction is in the adrenal medulla, as indicated in Chap 26 Similarly, the juxtaglomerular apparatus of the kidney and the islets of Langerhans of the pancreas function as neuroendocrine transducers, in that they convert a neural stimulus (in these cases adrenergic) to an endocrine secretion (renin from the kidney and glucagon and insulin from the islet cells)
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Role of the Hypothalamus in Sexual Development (See also Chap 28)
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The hypothalamus also plays a critical role in the development of human sexuality and its expression, a theme elaborated in the next chapter The suprachiasmatic nucleus and the number of neurons it contains is considerably larger in men than in women, a dimorphism that becomes evident during postnatal development The interstitial nucleus of the hypothalamus is reportedly smaller in the homosexual male, evidence perhaps that homosexuality has a recognizable morphologic basis (Levay) This biological evidence has been sharply challenged (Byne) These issues are addressed further on page 503 The intimate relationship of the hypothalamus with the development of sexual characteristics at all stages of life is shown by the appearance, in the infundibular area, of hypertrophic neurons that are rich in estrogen receptors; it has been proposed that some of the symptoms of menarche are timed and mediated by these hypothalamic neurons With aging, and more so in Alzheimer disease, the neuronal population in this region decreases markedly; the sleep disturbances of senescence and some aspects of the sundowning syndrome have also been attributed to this cell loss
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