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    Unit 4 Neuroendocrine System

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    The Neuroendocrine System Learning Objectives To be able to: 1, Distinguish between glands, endocrineshormones and target cells, and explain their modes of action. 2, Summatize the main endocrine glands and hormones. 2 Discuss the role of hormones in sexual differentiation and activation. 4, Explain the role that hormones play in the stress reaction. 5, Outline the main sources of interaction of endocrine and immune sys- tems and discuss their effect on illness. | fones-of the Bop) The neuroendocrine system consists of: (a B (b) the glands/ st Stee a smal po logical-eifects. In our dis “iin, first the basic principles of the néUFoendocrine system afe~covered: and then ‘he principal glands and hormones are summarized. After this introduction, the hormonal ‘fects in Three important areas are presented:.(a) sexual and gender development; ©) stress Teactions; and (c) health, focusing 6n hormonal effects on the immune system. tae usin lige le wilocrine;ayStems (e-f-those involved in heart regulation) tha: are responsible — ae ne marae heb ‘not discussed in detail; the focus is on those By anat have significant psychological effects) tony «°° 04 Of this chapter, i-should be posible to answer some fundamental ques neg. Piological psychology. (a) Why are we male or fei¥ale? (BY Is it possible for a (XY) male to develop into a feinale? (c) Do hormones acti vate our sexual desires bodies when WE faced by stresoy za vic disorders (6B, tpn," rappens it L and behaviour? (4) What oe PO a ime)? (2,09 ROT son the immUne Scand ten onal es” at exami a (f) What are the fey thor 0 cl se? o 470. Foundations fo contact with nearly all cells that is, those mje The lo Pormamitters op postsynaptic receptors (chapter 4 anagonseactone (hibing the Cares J: i but not ) exocrine glands ss; mostly on the surfacee! ‘which release tet this reason, organs wine imes called fowever, other organs (e.g., stomach, release hormones into the bloods n (but this not their primary pose); these rit ally all hormones are Feeasetin pulss 08 are part of the = (igure 6.2). Low minutes to hours), or but ident (most Reing, those hormones associate the menstrual cycle) Types of hormones Paracrine Blood system aw © & — Figure 6.1. Three main types of hormone-recept fF action, Ut) NOU eea ne BAC sical Factor’ hormone hormones are cited todnguish them from hormones that have been chemically defined), Most hormones Tall into thrge major classes (a) Amino acidderived hormones. olecules; there are two types: Peptidehormonesand protginshommones hormones,longer fat molecule; e. medulla) (b) ide hormones h: action: “none receptors fall mito TWO broad types: (odesuface receptors; and (b) intra Slice receptors nth mode of hormone molecule-receptor binding, dl the, Cll Activation of the receptor leads to the pro * Fist messenger is th Hormone lene in the blood: ‘Tephysloia ee ermine Blood concent: ula other Fe e “ene ee gulatet tyr, o/secreting gland eile instruct the producing ng, gland factate activity (POSTE feedbacks Tt the hofmone to drop by rah for longer and thus have lon; erlasting Agonist and antagonist hormones In the same way as neurotransmitters, hormones can act jer agonists OF sitagonists are molecules that bind to the rec ate events, nblock the binding of agonists (ig., blo yechanis ie molecular(hiaraonal) keys will not work). Many hormones: i, exerting sd el ample sin and eager hae opporie cf rete = insulin Tow evel: TE seth ¥ a ‘sechmnnsekcecergetpnne = ack" ithieved. Hormone-neurotransmitter differences poop ones lod vesseloin. specific brain regions). Hormon. i . action, much broader Wie elec ran RCUEE! neurotransmitter, which involves chemical ss the enapsicet see chapters 3 and 4) Compared a _pogrease: in space (tiebody) snd tiie ‘Therefore, hormones scrvg re-only.as a means GF gehieral transmission-oF information; 1héy cannot Saderspeticinformatn addi- tion, hormones often affect Gtiplessites"of action, unlike ancurniransiniter, which smeeracts (7a ani regulate s Body, sich as grow oT determining the envi : d esleg., fleeing from a predator; preparing the sex organs for teproduction; conserving blood glucose £0 maintain effective brain functioning), —— Although important differences exist Between the mode of release arfacion, neuro- transmit rs and.bormones share a number of common feature. Some hormones are local hormones that are released igfo regional blood vessels in.the brain, influencing neu. ich the same way ‘aporeamcin (Deutch & Roth, 1999), rene and neurotransmitters also breaks down when cor ‘ nical te.g., adrenaline and noradre- naline) that serve hough hormones may not see! In gefieral terms, itis possible to distinguish horn ance of travel. A chemi en UASK Yoursenr tance from 1 (b) whenituses the blood.as med [What are the clear-cut means _of contrast, neurotransmitters (a) are | distinctions ‘between the released by a mi ils.) | Stlons of hormones and and (b) have a WelFdefined onset and end of their action | is servin, @) Neuroendocrine Glands The position of the glands in the body and their major functions are shown in figure 63 Pituitary gland The pituitary gland is attached to the base-of the hypothalamus by a thin stalk that con: —“GiiSeurons, blood e. (Sek figure 6:1.) Thispeasizedseruc: ‘ure consists of, (@) the anterior pituitary, w arctan th scoz tute», which regulates the gins, emphasizing anterior. pituitary.is and (6) the pos EE! Te ewtolts _—— Pineal Hypothalamus ae ituitary Sea iiyrod Parathyroid Figure 6.3 Endocrine glands: position of the glands in the body and their major functin, x 4 yh 4 a ae ak Cos hypothalamus, a SS CaXr "i & Posterior pituitary Anterior pituitary sete Figure 6.4 Physiology of the paraventricular nucleus (PVN) of the hypothalamus, 8h! cell bodies of corticotropin-releasing factor (CRF) are located, MIS Vetere Eas Frontal Corpus Hypothalamus a callosum Optic chiasma ~ Anterior pituitary | Posterior pituitary Pituitary ae “Cerebellum Figure 6.5 The position of the hypothalamus and pituitary gland. Organ Hormone SERRE HEE EEE ‘panlanus Varios releasing hormones Stults ons lease ol various Homes by play Aero ptstary Thyroi-eimulainghamone (TSH) Stimulate hid Ltenzing hormone (i) Increases product progesterone (me) and testosterone Functions (mal) silat Folie stimulating hormone (FSH) Increases producto! oesoQen and maturation of ovum (tmal) and sperm production ae). acTH Increases seretn soe hormones by areal lan. Pralaein Increase mi Bender Pans rosue Poster piitary Oxytocin Conta ine onvoctons mi elese cea epoch parental behav and eoual pleasure “f Const ood wssliestlcos presaxe. Faure 6 The hormones and widespread functions of the hypothalamus and the pituitary sland, ‘ones fro $3€cretions control the tion the thyroid, a functions, iyseeretes@Totmone that tells te kidneys to decrease the amountnfwenee excreted in HEGrine, a useful me¢hanism when the ody is short of water (figure 6, The posterior pitafary is compo: I 1ypothalamic neurons that exten. ‘nnge bundle behind théantérior pituitary), the appe 5 5nthe hypothalamus fynthes Ofvarious other peptides, Hypoths ons to The hypothalamus"¥égulat feurons in, plus smaller amounts tary = including, or for-each hormone. vin the release of thes sing fac wo tyoith oh nal HEFE Pal tn, ead development ‘attached to the thyroid are foury, release by ich og Ta which binds to ane, 3 imulates the antefionspituitanyato sr, hae blood to cells in the thyroid gy cavels in U — a5) which affect (mabe rd Boones ha ces this inhib eee hus reduces therelease OF the roid (HPT) axis is an example of the action in themselves: “he hypothal mus-pituitary x a mega fedback, mecha W 1 then fll bx is ian a 4 Tee Pancreas “The pancreas is located nest, all intestine; and (b) it discussion is concerned only with the s (there are app" “The pancreas is domposed of many s mately one milion sets in human being), Pancresir lets hhouse.three major cel f° aach of which produces a different hormone: oa els serete" (eich the ges Tees ‘oF incrcasingpblood gHICOSETERST ing sufficient glucose is available for the brain) (bells produce ini a4 @ 7 oH involved in growth and development) ri a juctionrof insulin lm secon Adrenal gland (rhese triangularshaped gfands are I are impomant in me axis. The adrenal gland is adrenaline and noradh UT ils SCTE 2 nd _co functionally separate and Fave different embryological ovigins, , ee rhe adrenal medulla synthesizes and secretes adrenaline and | noradrenaline, which have the same effects a8 release {rom the neuron terminals of the sympathetic branch of the autonomic nervous system, although the effects of these hormones are more long last. ing. The physiological effects ofthe secretic by the receptors 70 Essentially their physiological won with dealing with str nstriction of blood | vesels {hefeasing blood pressure increased metabolic rate; dilatation of pupils cessation of non-essential processes (eg, digestion and motor activity); arid increased sugar out pucoftheliver ihe adrenal cortex isa site for the manufacture of many. steroid hormones. includin, sex hormones, especially androgens; the production of egiticosteroids is especially important (see below), fineal gland (also produced elsewhere in the body: e.g. inthe Guo at right esata Se cas eee nin promotes deeds cue oie ava ae id adrenal cortex. caeetee Sex glands: ovaries and testes ‘The testes and ovaries serve two functions: ( amass) and flies (nies) g acteristics and in saviour; sro hormone ogeterne preps the uterus for inpl ale tess are responsible for the-pro- duction. ofese n adrenal gland), which, 5 tur, is respofsible for male cfaracteristics aid"Sewal’ | ASK YOURSELE “Abu The action of Use two hormones is discussed in more | 1 relavon to the cenval ‘ail belo In the male foetus, the gonads differentiate imo NeN°%8 ee tation, and : 4 es rater ‘gure 6.7) u Neuroendocrine Hormones This lo, gee tO8 Summarizes the action of the major hormones; then in the sections to fol: HPSACtion of some of the more important hormones are discussed in further detail, ee z d qu ¢ _- urethral fold i : urethral slit 2 — Tail (out-of) 4 : Ad y ~ Anus — : seventh to eighth week eet! meatus Twelth week Figure 67 Wolffian and Milleian systems and development of genitalia. At six weeks the hus! Figure he bases of Woalfian (male) and Mallerian (female) reproductive dt hiss indifferent, undifferentiated, stage (2). From the seventh to eighth ws x the inf wf testicular testosterone, the male Wolffian system-develops,,and the ‘nhibiting substance cBUSeSthe-Miillerian system to des he-Mllerian syst generate,)in the absence thes Molen rte deveons to reproductive diets andthe male Woltfian ste" “t 7 the female and rile reproductive systems are GeV" ¢ Insulin depe exclus fe of glucose, ae a lin release. However, the mer¢“ight or smell fed ae leads x r pancreatic insuliyfsystem allows the body t maintain constant BIO concentrations ea ose despi Seer ryibg Frequancy-and’c coneeneof meals ~ without this system, skipping breakiast could fead ta, coma and death! ae = Adrenocorticotropic hormone (ACTH) Adrenocorticotropic hormone (ACTH) is secreted by the response to corticorropin-relea pothalamus; CRESis=released Yin ‘Spon fo eon Oro ct elon). ACTH ao cl —— the adrcucasateegcng to the Se is itself glucocorticoids, which ac ‘The release of, inner sin low, this system mediates long-term stress, whi can lead to a number of diseases. ‘Among its other functions, ACTH is invlvedimthelexpectatioWthawsleeprcomes:to an eae cre is a marked increase in the concentration of this hormone one hour before awakening (Born et al., 1999). Thus, ACTH appeats'to"arouse'the body fomaction — this release may occur too early in depressed patients, who often experience catly waking. ‘common test ofthe response of the adrenaligland W6"™MGTHLis the “dexamethasone SuppFEION test’, in which the : paisley which issaisyntheticssteroid ve is given and levels of cortisol are measured. Cortisol levels should response to the administration of this chemical hich in normal individuals i lease of - land and thus st e secretion of cortisol. This test identifies CE) Ggettnteetes to,a single ob: the ach re ig Respomse-t aren rena RHI ea fom the a) =e hormone (ACTH; see aboye) from Arginine vasopressin Arginine vasopressin (AVP), product sed by the posterior pituitary, et | Too cis role, AVP is often a an rth, ‘ADH)/AS a -d : oben “easement ata dny terminalyat the posterionpituitary gla jored. When the bo sa intr datl by neurrs wh en i s them: -nto.the Blood, which tne of AVP conta newony Ces 1 vy itto the kidheys, where it inh ‘ring. When there is x In addition in pair bon e Oxytocin Oxytocin is produced by neurosecretory"€élls"oPtheyhypothalamus and transpeny to the posterior pituitaty gland (inthe same manner as AVP); itis also secreted wits on responsible for: ejection of milk fiom ty breast; stimulation of muscle contraction at birth (itis released during childbirth vu the foetus stimulates the cervix and vagina, enhancing the contraction of the musa} and ivestablishes Wate Behaviour (e.g, its injection into virgin or non-pregnant rt leads to maternal behaviour; antagonizing the action of the hormone leads to a flr of the mother to accept her pups). The body releases'enormoustamounts of oxjtatt during orgasm (3 to 5 times the normal level in the blood), in both males and fas leading to the ‘afterglow’ of orgasm. Like vasopressin, it is important in pair bona and attachment. ‘Oxytocin is sensitive to touchan/ physical contact, and is one ofthe hormones imilt! in romantic Joyestt can be thought of as an unconditionedstimulus (CS), which forces the conditioned Stills (CS; e.g,,a particular person) associated with its ‘There is a cycle established such that the release of oxyt oh (Sippbihettreentantem ching ‘This happens in pair bonding between m=” and baby, as well a between lovers In the case of sexual behaviour, the release ofaxyteciiicTeases sexual desire and ivity; it leadsto the incteasein testosterone, which increases sex drive in males and f™!® In females, i tirmulates the release of oestrogen and prepares the vagina for pene ‘Oxytocin also increases the sensitivity of the penis and nipples and makes ongast™8 further stimulation ofthese organs then leads to a further increase. ‘The effects of oxytocin are la srs of 5 nse 8 cess JHA hsreson sem hat les iH Se ne eT ~ Luteinizing hormone Blood levels of hormones “= Follcle-stimutating hormone 1, rogesterone ' 5 10 45) 20 25 28 Days. Ste at /°Folicuar + ~ Lute phase ‘Menstruation Phase and this sensitivity ier over the menstrual cycle, with greatest sens: ce ovation. tao appeasharaRoT pas eaonars ual activity aier-an argument might redice thinking about 2 prior argui “ust one to forget about some of the distress caused. Experimental studies support the role of oxytocin in sexual aviour. Onytocin (a) ri ) in virgin abindvces full maternal behaviour Within minutes f injection, and (o injection of an “Stocin antagonist 1 effects of oxytocin) suppresses the i Tal beh, nT rever, these effects are not seen after several days Of maternal coxytocin is involved it the onset rather than the ian, Mayes & Cohen, 2002} Tanga! behaviour, indicating that ~~ Somatotrophin and somatostatin J and secreted wth hormone (GH) ts synthe c _canterionpituitary. The major fianetion of thie gowth, Pr ‘ ei oe gamma) . Melatonin ‘This hormone is released by the pinéaligland/andis controlle tq sqgssgMElaronin secretion iyed to the thin. Passengers any across time zones often take Melatonin tal Ge. t0 phase-shift thei Ti to match the night/day phase ofthe eye, mh ve ——— Tine “i beaten Geen eae (SAD; a type een is in. [northern hemisphere count oe "year, the pr PrevalencevOF SAD 'RIGHSAD patients are which consists of SI the 7 es : Gonadotrophins This class of hormone Fhe ae eRE DUD pen ‘ fori, the gonads hibits further synthesis and seri oft “Tae HoPwORE LYS both se ; oth ems LH simulates the seins (eststerone from the testes; oestroge 3c ovaies). Fal deptecrnads pater estrogen from the oval ge quantities of Lt mature inthe" ovaries during the preovulatory phase of these follicles travel down the fallopian tl fertilized egg jérbedded in the wal o the hormone (FSH stiles the (natraion a for sperm proxhietion——~ i ir ina wterus (ic, ce womb). folly so of ovarian Yollices; and it i aso "°F Hypothalamus | GnRH T + Anterior pituitary —l FSH and LH 1 Gonads (secrete sex hormones) -{_ Sex hormones Reproductive tract and other organs, Figure 6.9 | Hypothalamus-pituitary-gonads system. Gonadotrophins stimulate the testes and ovaries and the pituitary. Gonadotrophins comprise lutelniang Iernen ile stimulating hormone (FSH, the cegulatar ot whieh is secretion of gonadotrophin eeceieg hormone (GnRH) from the hypothalamus. Negative feedback from the gonsdx EIBIis Te he, synthesis and secretion of LH and FSH. Prolactin ‘This hormone is secreted by. by other cells in the body, production of, mammals (i.¢., lactation) and stimulates th ovaries to secrete the hort 8 flcts go beyond these functions. For exam] and secrete prolactin, sugge Sr pituitary): when this hypothala and secreted Beasties Sex hormones The gonads do not just create sperm and egg c ate teas ora. ‘two main classes of gonadal hormon estos a iol te most common oestrogen. 1H, ovaries produice boll Horm st compared ya ith women produce @ trifingsamountoftestosteroneiust enough for norms) ons although it is sufficient to enhance sexual ACuVIEy); a fo 29 (approximately 7 mg per day). These steroid hormoneFAREaso Snipe dy released from . Androgens are called ‘maleshormones"*because their le ch higher in than in women. Oestrogens-aré often referred to as - ad level i ‘The ovaries and testes release a class of pe, clean cor of which is'progesterone, which in females pe themierurteg, by decreasin ind breasts for pea In Tales. gerogenispasent in low SORRENTO in the blood, but in very. » fentration in semen (in fact higher than the serum Concentration in fernales) has led to the suggestion that this hormone plays an important part in rate fe etal, 1997) = By i Testosterone ‘Testosterone is secreted in the bodies of both males and females, It is scereted fume le; in feral crest y other hormones mnadotrophins) th ted into the bone stream at the pit HY gland. ‘Testosterone core eae al behaviour and seems to besimportant i"domitance:hierarchies (e.g, defeat of mk Primate leads to a decline in this Kormone o sway ch i dominance ~ this might be the sensible short-term strategy given Previous defeat). In the case o¥ sexual behaviour, testosterone levels are highest berwet? » When sexual motivation is at its. typical peak. “ estrogens steroid hormones, and as such ener is comtroll& ee *ex than in the other, For example, oestroge? oe Sponsib developmentin women, and androgens activate the § " peponsbl forthe growth of facia hsicin men. arafoporm sacs he gown hair in both sexes. Thus, the XY differences kez> noes St Hatt Y counts den mean aes ae cfs ditferenfeeet oF androgens andoestrogen his were not the case, thei hei would be the same in both Sex cs (albeit to varying degrees, depending on he“ centrations in the blood). It is therefore not arn ; aeswetn i that bind tg fntracellular : tion see above’ o After removal of the testes o, i : tines in Pe . viourt declines it tion to thé blood concentration-oF the sex havioue deo esl hormones, Testosterone restores “ castrated males; oestrogen a oe™ ates sexual Peo seme —_— ecie: a a of species, AES Gompere-oracees.t. females sexual selection; s¢€ The Neuroendocrinelsystem) 185 2% ecess. Increasi ciated wth ‘nggpasedegerssion “Thus, sex hormanes_are fundamental to the key com- ponent of fitness: differential reproduction. In summarizing the major effects of hormones, it should be remembered that they work in a synergistic manner; it is too simplistic to associate one hormone with one effect or a few isolated effects: the action of one hormone often de pends upon the actions of other hormones, ay have multiple effects (Nature favours adap : that serve more thanyone purposevthis makes efficient u: § OF resources and allows newly evolved functions, to Build upon jre-adaptations; see ch ter 2). For example, corticotrophin- releasing factor (CRE), vasopressin, oxytocin and_ad) i cotropic hormone, (ACTH) all seera-t st fear_and ig 5 at extent are we anxiety, as well as on‘learning and memory (Croiset, Nijsen ;, ng and slaves to our hormones? & Kamphuis, 2000). ean The Menstrual Cycle The female reproductive system is characterized by its periodicity: the aestrous cycle, which 's ‘pically 28 days. This cycle is the outcome of the interactions of ovaries, anterior Pituitary and hypothalamus. The cycle starts with the secretion of gonadotropin-releasing hormone (GnRH) from neurons in the hypothalamus; the feedback effect of oestrogen at the anterior pituitary, and possibly-also the hypothalamnus, i assocaeed so ith excitatory and inhibitory effects (see figure 6.8, above), Poxftive feedback is see: re a surge in luteiniz- ing hormone (LHY precedigg_ovulation (ifthe male there is only negative feedback at the anterior pituita At the meron ‘menstrual cycle, the anterior pituitary releases follicle stimulating tormone (FSH), which promotes the growth of a follicle in the ovary; this follicle murturce the ovumt (gg cell) and leads v6 the production Sf oestrogen, Towards the middle of ‘the ‘cle; the foicle builds up more and more receptors t0 FSH; so even though the actual concentration of FSH in the blood is decreasing, its effects onthe follicle increase. As a result, the follicle produces increasing amounts of oe type of oestrogen, oestradiol. “This leads to increased release of FSH, as well as a sudden ‘Surge in the release of luteinizing hormone (1H) from_the anterior pituitary. FSH and LH “ause the follicle to release the mone progesterone, which Sp aT eimplantation of a fertilized Quel Progesterone also inbibits the further(release of LH, At the end of the menstrual Ole The level of LH and FSH, oestradiol anc progesterone all decline) If the ovum is fenilized then the level of oestradiol and progesterone increase gradually throughout Pregnancy; if the egg is not fertilized, the lining of the uterus is cast off (through men- Smation) and the cycle begins again. os mtraceptive and birth pills work by disrupting the feedback cycle, between the ov: inathepituitary. The combination pl contains both.oestiagen anid progesterone, caries it works pein Ways: high levels of oestrogen beginning shortly after the end. of the eran! The efSUPPTCS the release af FSH, thereby preventing the development of the follicle wal lease of y e hes tha the ovum, Progesterone blocks the scctction of LH, thus further ensutin ‘De ovum is not released, Unlike women who show increas l sexual fantasizing, mid-cycle (the periovulatory - shaviour a sexual ce tion-in-the blood and>sreig po e050 SOP reac at tis period (Adams ota vy is important IP maintaining sexual motiva en 8 __ % j_abeir menstrual cycles (Mety behaviour 7 ee fee, “von wh een Fe co fl se ms oe 999; Weller, 1998); the OS 3 Ei th, dee Tike most othe! ~ physiology . ing human beings Cn a si ea tig chennieab signals. je for this i hE ey, To test ‘whether pheromones. ‘of donor women} a wyatt, 2003). To tes othe underarm cecretions of donor women} Most jonor’s (Preti et al., 1986; sour women. suet SPO eae 4d their cycles eee wom he ink (998 da Si reer ahe armpits of WOMED. taken at diferent phases of ther les comp less of LL selene 98 eayed depending on doen cyele, affected the release OSS one sexual traction. The effects of pheromones dp, ‘itost mammals use pheromones on activity of the gomeronasal ons (a set of receptors Jocated near the olfactory rey cars, Ie is & prominent structure in. OS, ‘mammals, and is ess cen th foetus, but in adult humans ‘cis very small (Mont: Bloch et al., 1994). There is, yx Te exidence that pherot .man beings in sexu: | attraction Howe little evidence that pheromones are used [in human belngs in SOA = underarm secretion, when ple lide eis do gaggst.a sexual 0. For example, male URES ST (for sir hours and masked by fragrance) w females alters the Tength and timing ¢ er Prenstrual cle and affects mood increasing_ relaxation d_reducing tens aaa my impact pon several components of fertlfynendocrine responses (prin ects), information (signallereffecs) or mood (mods effects), behasiour-teleaser jective. might ap ‘ffecs), But why should this phenomenon exist? An evols Shavit would be an advancage in funter-gatherer societies for women 10 be fertile ae! the same time in the presence of men, because groups of men spent a considerablep portion of time away Huntin : : Fr rmanals, the effec of pheromones ase_widespread, For example mt cepa constanly monitor the urine of their female hast to determine which ones are in “heat” ct a fer) ~ the vomeron ees e lu" phant’s mouth. In the urine is a pheromone that activa! onary advantages of5y%- chronized ovulation? hardwired circuit in the brain that leads to an insane 3, 1996). TN tion (Rasmussen 1 L] Sex and Gender / gan Hormones play a fundament () during aber inthe eta ole (a) in the foetal organization of the, se8 breast enlargement, voice breaking) fh tse seaual characteristics (8 PU ic of sexual desire, motiv: ey also play an importat in the ati motivation and behaviour during sduttheud, = cua jzational effects organ" Organizational effeets refer to permanent change in the structuréi.c., morphology) of both the sex organs and the central nervous system 4s a result of exposure to hormones, ‘These changes occur IT Tren! periods during gestation (irr tHE Ease OF human beings, about the thigd and fourth months)) during which time primitive sexual structures are ost sensitive to hormonal influence, Gonads” In all mammals during very early development, the gonads are identical in genetic male (XY) and female (XX) foetuses — this is the palgrenciared ‘sexually indifferent stage" io Differentiation isthe process of the formati the male or female reproductive sy=- tem from the undi iflerentiated structures: Miillerian ducts and Wolffian ducts (see figure 67 above). These primitive structures degenerate or develop to form male testes or female ova,' and the external get ‘hat determines the developmental route isthe pres- ence of testosterone: exposure to this hormone leads to development along the male line; in the absence of exposure to the hormone, the primitive ducts develop along the femalé Tine. “jhe mechanism by which testosterone is produced in XY males is the presence of the Y chromosome, which contains tas that, Tea the {0 thé ducts developing into early testes? the production of these testes then Teads to the production of resrnsterone and the fur ther growth of the te = 7 and thus to the production of more testosterone: this, js the process ef gonadogenesis Testosterone also causes t€ primitive Wolffian ducts to develop into seminal vesicles (sacs that store semen) and vas dderens(g duct Frofa the testes to the penis). ® peptide hormone, Mifillerian_inhibitin, tone_(MILD,feleased, from the-testes;causes the isllesian ducts to degenerate; time the t&tes secrete testo: Tone, Which ‘simetates the development of the Wolffian duct system to form the male sexual organs. _, In contrast, females develop @long a default developmental Path to form | ual genitalia made from the primitive(Millle Wal he female sex- her Wolffian duct degencrate and her primitive Miilleriay ducts develop into fernale structures (uter id vagina). Qvariai (oestrogen) hormones seem not to be involved in early sex- ual development; without the effects of testosterone (secreted by the testes) the default (xis female. > _ So ‘Thirner’s syndrome is found in women wit ‘only one X chromosome (so-called Xo; see chapter 2). These women do not develop ovaries and therefore adequate levels of estrogens, Which require two X chromosomes; however, they do develop female internal sexual ofgans ‘and-extérnal. genitalia. This syndrome seems to show that ‘oestrogens are Not required for female sex organs to develop, Therefore, it seems that androgens cause ‘ales: the absence of Sex hormones leads to the formation of the female phenotype > meen itis the production of testosterone during the: critical periods of develop: Uestonte esstermines maleness (presence of testosterone) and femaleness (ab “rone) in terms of genitalia, internal reproductive or gans and psychological ce of ey Gores jas development in favour of one of xy characteristics. X and Y chromasomes Pi = spe shown that. if genetically ae oy sions of se}, In experimental animal rial periods, then they develop a . sed cron€ during these £4 zi a tenons dt ih human beings natural experiments OCSUrin whighse al ‘ Viduals develop(both morphologically and psychologically. along ale and female Androgens (principally testosterone) pass the cell me mbsane of in cals ing neurons) and attach to receptors in the nucleus (as aq = He boson androgens are then, stradiol (the principal type o! — wich alters thé “expression ‘of genes within celle. By this means Benet eects ae ema to forma, orphid) physiological and psychological sexual characteristics, Tis ime, aaa ‘her oes cr serorerone edited by the scl eral to oestrogen. (The fact that oestrogen in females does not lead to this same outcgn, due to the présence of an inhibiting molecule, alphafetoproteatsee below.) ——™* After playing this fundamental role in the organization of the reproductive g inch the brain structures that mediate sex-typical motivation and behaviour, he efi of s&chormofies lie dormant until the period of puberty,when there is a furt NF inp, from(fonadal hormones to sexual development. Boys éxperience thelr voices "breakin and secondary sexual characteristics forming (e.g,, pubic hair); girls begin theirsaensiny cycle, gd fat is laid down on the thighs, buttocks and breasts. Puberty is initiated the hypothalartius, which releases luteinizing-releasing hormone (LRH) at a rate of one bm, per hour (these bursts seem to be triggered by(body weight). This releasing hormam stimulates the pituitary to secrete luteinizing hormone (LH) and follicle stimulating hormone (FSH), which in turn stimulatés the gonads to release oestradiol in females and ted? terone in males. —~ Sex-determining genes Genes on the X and Y chromosomes play a major role in sexual_differentiation. Two major genes arénvoTved in this process: the SRY male-making gene and the DAX1 female making gene. a. SRY male-making gene One clu¢ to sex-determining genes came from the study of XX men ~ that is, men vi! the female XX chromosémes (the first indication that they are genetically female oft* comes withthe discovery that they are sterile)) In these men, it appears that a tiny P*" of their father’s Y chromosome liad become attached to the X chromosome - it paternal X that is passed on to the son. A search was made for the male-determiaae gene on the Y chromosome, which was absent in XX men) This research reveal = base pair sequence.—a viny gene christened the SRY (see Cookson, 1994) and 12% on the short arm (top half) of the ¥ chromosome) Around the sixth week of fost i'ig this, gene that causes the undifferentiated tissue to develop into testes’~ without! activation, (the tissue develops along default female fings, Ie is remarkable that ? with so few base pairs (many genes are composed of, offen many, thousands of bast could have such a profound effect On sex determination) UE eee neu An XY offspring may have the SRY gene missing fram. the Y chromosome (or the gene be inactive), hence this chromosomally X¥ boy devilops into a girl. In all respects he chil appears to be a girl, struation does not occur, and then the problem is identified. pax1 female-making gene ‘The absence of the SRY gene, or its inact leads to the default development of female genital iD Another gene is important tooPAX. is located ¢ of the X chromosome» and acts at the same point in the“dev ental_pathway as SRY in normal males SRY_wins outcand Sexual tissue develops along male lings. However, if ‘the DAXI gene (enter the individual who is genetically male (i.e., XX) to develop PhysKeRT as a females seems that the fanctiop of DAXI is to ensure that secondary m@t€ gehes are turned of t@ anGnuimale gene) in tog strong a dose (due to duplica- os the SRY gghe and turns off male-making processes, The ‘female’ often finds out"the truth when they fail o menstruate at puberty) There are also chromosomal XX females whos ‘velop as males/ Because of the activation of SRY and the inaction of DAX>> This, SRY and DAX(intetactions can _account for the ‘genetic basis of sex. reversal syndromesemtechnically kn: 1” (Swain et al., 1998; also see Meeks, Weiss & Jameson, 2003). ‘The central nervous system In addition ro the organizational effects af early exposure to testosterone on the devel opment of the gonads and sexual reproductive anatomy, the central nervous system also - a ectvons syste undergoes offanization along seXspecfic ine=The most obvious effect of this SeCspecitic organization js on the(hypothalamuswhich controls the anterior pituitary, the master sland), which, in the feyfale, is responsible for cyclic patterns of hormone release from > the menstrual cycle). The male hypothalamus is incapable puberty to menopause (D Chsen is cyclic release. of hormonésSthe same is also ‘genetic (XX) females Sparel fo High levels of Conon their Gritical period of development) who have developed along male lines. — ce Ras An example of thi{ hormone-related sexual dimorphism is found in the medial pre (pric area (MPORY of the hypothalamis. Fhe MPOA ~ asexually dimoyphie milous ~ is {Pally larger in males than females, and its stimulation in males causes male-typicit laviour, af males, female-typical behaviour. Destruction of the MPOA results inimpaiy€d sexual motivation’ inmost species, Dopariine and noradrenaline are import- ‘yGsurotransmitters in sexual behaviour’ Neurons in the MPOA increase the release gyPibine during sexual behaviour; sexual behaviour also increases dopamine release 'theucleus accumbens, An important structure in the brain involved in ceintorcement snd erate "xposuie to androgens i rattlnization. In the rat, xf te (geay itch & Denenberg, 1998), that is, masculiaiZation a play mechan tn ‘s ‘hat inher ipsensitivity to androgens show asfemale-like pattern of play. the sexually developing foetus causes a proce sure to testosterone sensitizes the male-rat vo may a Pein : feminization Maseulinization and f ajong evo axes inthe Brain: (2) maa is along, tWO axes 1p ENE DY a EEATTOR TT : até testosterone has tw@ effects on the central q é ological cifeentiati z ‘SyemmmzaTion The ma 7 oy, izing ( ding to the male phenoy a, : jeminizing (corresponding to aS onan cnt oo ‘effects occur: (a) demasc' ew) Fei i Ineo dhs female phenotype). These two axes pros luce four possbieapy (corresponding to i Ene cinivisation (typical male development), &) me oe ie oo (typical female development); 2 Seannizanon = applal nak de ulinization-defeminization (atypear Catypical went) and (d) demasc i. ae fan Oeggeaens ‘may be involved in the femiaiston of female being in developmen \Fitch & Denenberg, 1998). The development ot ete : Pon exposure testosterone at diferent stages of foetal development. nCae oF testosterone are siediated by the oestrogen oestradiol, which masculinizing effects of sexual behaviour GPis thus somewhat misleading to talk, Of ces, eas a ‘female hormone’). In the female, testosterone (which is present in low og ia pha- fetoprotein whi centrations) does not have thi¢fasculinizing effect because alpha it Which ng present i se eS ese nn blocks ixfrom Tering hc ieeg ee enteting the cells that are developing during thé critical period: if this blocking effeas, not happen netimes it does nor) then oestrogen. pfoduced in fernales woul, tormsciinization (in rare clinical cases, this protein is defective) leading to titres. izavionof the female's brain). The precise effects of oestradiol depend upon the stifey development. behaviour; fefhales exposed to androgens during early development form maleypr behaviours. A mascUinized, but not defeminized sar would develop a strong pret w both fo » mounting and adopting the-femalé Rt i pei roc nt © sees process underlying favaaes male homosexi, Pa iu is known at present to allow frm cons i Psychological differences also develop di por aoe monkeys ex; sed etsierone during their sensitive Period engage in-more 10 Play tian other females? y5e more ape maa id ; “a essive: threacenif es (Goy, Bercovitch 8 Mera ee eno MERE OF testosterone-Gy show red typical sexual bets luced female-typical sexual < Congenital adrenal hyperplasia, CAH?) = boys’ toys’, such as cars, and wer : seems that “Thi i "ed playmate (B, & Hines, 197) behaviour is he beeen between Prenatdl ance. ae cl gender est- male and oestrogens in the female are secreted intg the general bloodstream and exert \2Fus effects throughout the body — During foetal life and shortly thereafter, secretion of GnRH, FSH and LH and the sex Sormones is high, corresponding tothe period of dillerentiation, Ai puberty there is once ‘Bihashanp rise in atiyty in this axis-Phis reaches a stable level in males, but in females there ig Oscillation in the axis corresponding to the menstrual cycle (there are changes irezesponsiveness of the anteHOr pituitary to GaRH and Of the,ovaries to FSH and igi the aay period In both sexes, impairment ofthe pituitary results in a loss srl dive; sex irve tends to be reducectin-men with a reduction in testosterone >> coy He dominant effect of testosterone 6p sexual motivation in the male, there is Somparable domi ifat hormone in the female; both oestrogen and androgens ate a SU aiaoue in various regions of the brain ceptors in vaiou Ne rn ised Oogeogets Hn fala and septum). Ns in matey, i = yal an septum). Sy sexual arousal pola Se vain non mm ee le sex " Pineal role in hum h seats in the os" FETA a a androgens are produced i both xt inthe wi from thea 5p The role of androgens in fey i they a ale ed in the teste3) The.rol in Fema males they aes rie n do not cease afr the mens, wy sexual desi an fer the ms oe Er nc, Ament oF (ich pg per cent of oestrogen) sige loss. ofits” FSMijand LE and thus» we onan release OF OEStFORET : ae oa destrogen ave eflts inthe bn Jated to sexual behaviour. For example, they increase production of, **y and werotonin rocetors inthe nucleus secs, the prefrontal corer angyye® oy cores these reefoR APE TNR To Be associated with emotion, reinforce ‘mood decine in oestrogen ould therefore, eed w(decrease activity YY ese) tors and thus emotional canine declingin oestrogen just before mensiniy afer giving bith and during the mengpause tay Be response for mood chang seen during these periods. Organizational effects vs. activation effects Jn discussing the organizational eects and actiaton errs of hormones, it was noteiy the former/fvolve long-term structural changes while the latter involve short-tem ay von of esmatitn and chavs: However, the distinction between these py pes of 6 not complete, and should be thought of as one of degiee Fthertin kind. For example, neural systems are plastic (Gee chapter 7) and subject 16 thy activating hormones - sudi neural systems are said to We ‘permaneiitly transien? & Denenberg, 1998). Also, sexual organs can undergo structural changes duriig sit hood. For example bod) lers in search of the jerk steroids ae liable ve develop smalter testes and sometimes experience breast enlargement Wises tural effects are duet0°a negative feedback mechanism that instructs the bran nt stimulate the testes to produce téstosterone. The brain, in particular, might Bethe a5 2 Semi permanent structure that is undergoing. confinuous restructuring. 7 Sexual orientation Sexual orientation concerns the relationship between ali and moxvatior heterosexuality fas a hormonal basis, then does homosexuality? There docs not to bea testosterone difference in heterosexual and homosexual malesalthough the some evidence for greater testosterone in homosexual femalé&-FTéwever, 28 sho"! ‘mones exer Complex elfecs and their effects interaet wit host of psycholog 87 ‘There is sonie evidence thatthe thid.itersttiahmucieus of te anterior POU §s smaller ip homosexual males)jtis also smaller in females {irrespective of 0 ental (LeVay, 1991) 80, the suprachiasmatic nucleus homosexual men cémmpared with heterosex: sng aus is 1.7 tres ‘of the hy ete 1 ames BE tal ones (cell numbers are 2 Ir ib & Hofiman, 1990). However, fo difference in this structure is found between the Suggbuch research is still at an early stage of development, and it will he necessary to Searthe outcome of this work before firm conclusions are waible. However, it is conceivable that sexual orientation 1s poveult of(iypical_masculinization_and feminization pro. cesses dosing bay deve nent — if this theory is true, then jomosexuality is just as ‘natural’ as heterosexuality. jour ever escape constraints? [LI Sex’ Development Abnormalities , Iris known that chromosomal sex (i.e., XX vs. XY) does not determine phenotypal gender (although it does bias it yery strongly in one direction) —‘the path oF trae gender des not always run smoothly. Without the release of specific hormanes al.critical stages in development, we can find chromosomal ex and phenotypal gender to be different. These developmental abnormalities throw important light on the nature of hormonal influ- ences on the organization and activation of sexual behaviour. Intersex Testosterone masculinizes the formation of the genitals and the hypothalamus during sensitive periods of development. Exposure ofmales and females to too much or too, intle testosterone can lead to partial masculinization of the genetic female (e.g., over- production of testosterone from the adrenal gland) and feminization -of the-male (relative insensitivity to testosterone). Various atypical hormonal environments can lead wo terphreiin uaphroditism, ¥,¢7, tndWieKials whose genitals do not correspond to genetic séxJn rare nstames, Both female and male genitals may be present (true hermaphroditism): one gonad niay be a testis, che other an ovary.datéral hermaphroditis)>or one or both gonads may be a combination of the two (an ovotestis). More commonly, dev is inter. mediate between fall male and female phenotype (pseudohermaphroditism; or intersex) ‘These eHfects are obvious in the of morphology (e.g., external genitalia); they may be less obvious in the case OF, sychological intersexism, but no less important. A num- ber of hormonal-relat ions have been well désctibed in the clinical literature. Androgen insensitivity syndrome Some women who have (experienced difficulty in conceiving receive the shock of their lives When they go Tor-a-medical examination. It sometimes (albeit rarely) turns out that women who §n all outward appearance and in motivation and behaviour are female are, fcr, chromosomally male (XY). genetic mutation prevents the development of functioning androgen receptors, and-this prevents the masculinization of the hody and nd there is a-defect in the androgen receptot gene on the X chromosome (this is rors prevents the deve, Prieta drogen receP! still has its gat an on sence of fun infllenee Lies c absence Sever 1 (WEEP flop. rogens, but th mA afin ale sx ol ingeflet, wi REFN a ete) fai t6 form (the VASE ae ‘second disonden fesistem ‘Mllerian inhibiting se er ei ic rant with bh sets of intern is ment Delayed penis develop’ : : | dition sen in the Dominican Republic. This gen ment ly balls at twelve), prevents the grow g Sai forms of masculinization); childy a a Delayed penis develops condition, known & gueveloce ( pe 2 ale) What happens a puberty when they Brow apy this dsonfer ar ra u if a female identity? The ansirer jr socialization Iead to the development of a. : Wold thei icc is plain: in most cases, male identity is adopted ay, heterodialoninaton {although a smaller number of individuals prefer to remain the female role). Damage to penis ‘Accidental removal of the penis offers another instance that allows us to examine the nature-nurture issue of sexual identity, and in particular the gender neutrality hypothesis, which contends that before te age of 3 years gender identity is not fixed, but affer a age gender is locked i. Circumcision (ie, the removal of the male baby’s Eri) sometimes does not go according to plan. In one high-profile case, t00 high a level of electricity was.used to remove the foreskin, with the result that it damaged the peris beyond repair. The interesting aspect ofthis unfortunate eve, in P te_event was that this baby wa! one of monozygotic (dential) ait The physicians an, oe never!’ and ‘I would rather ‘cman eek Dovid remained depresed rears old, he drove to a a "104 parking lot, placed 4 said, id,

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