Glucocorticoids. Introduction to steroids in general: synthesis, regulation, mechanisms of action, development, physiology and related - diseases
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1 Glucocorticoids 21 Introduction to steroids in general: synthesis, regulation, mechanisms of action, development, physiology and related - diseases Adrenal gland: anatomy, histology, hormones, effects, feedback. cortisol secretion, receptors, mechanism of action and effects Cortisol: effects on glucose metabolism, anti-inflamatory action, regulation, synthetic cortisol, the stress response Cortisol related pathologies: Addison's disease and Cushing's syndrome HPA axis Introduction HPA axis and its story lines Page 1
2 teroid and enzymes VLDL cholesterol CC pregnenolone 17alpha 3BD hydroxylase progesterone I 21BHL 11-deoxycorti... 11BHL corticosterone 18hydrxylase 18-hydroxycorti... 18dehydrogen... mevalonic acid AcCoA pyruvate HMG-CoA reductase 17 a-hydroxypreg... DHA 3BDI 17alph desmolase a 3BDI 17 a-progesterone androstenedione 21BHL 11 - deoxycortisol ßhydroxysteroid dehydrogenase 11BHL testosterone (*) aromatase cortisol glucocorticoid estradiol estrogen / androgen aldosterone mineralocorticoid (*) metyperone test Adrenal and gonadal steroids have cholesterol as their common origin teroid and enzymes gonadal steroids VLDL cholesterol mevalonic acid AcCoA pyruvate HMG-CoA reductase CC pregnenolone 17 a-hydroxypreg... DHA 17alpha 17alph 3BD hydroxylase 3BDI desmolase a I 3BDI progesterone 17 a-progesterone androstenedione 21BHL 21BHL ßhydroxysteroid dehydrogenase 11-deoxycorti deoxycortisol 11BHL 11BHL testosterone (*) aromatase corticosterone cortisol 18hydrxylase estradiol 18-hydroxycorti... glucocorticoid estrogen / androgen 18dehydrogen... mineralocorticoid aldosterone (*) metyperone test mineralocorticoid glucocorticoid Adrenal and gonadal steroids have cholesterol as their common origin Page 2
3 teroid and enzymes glucocorticoids mineralocorticoids VLDL cholesterol mevalonic acid AcCoA pyruvate HMG-CoA reductase CC pregnenolone 17 a-hydroxypreg... DHA 17alpha 17alph 3BD hydroxylase 3BDI desmolase a I 3BDI progesterone 17 a-progesterone androstenedione 21BHL 21BHL ßhydroxysteroid dehydrogenase 11-deoxycorti deoxycortisol 11BHL 11BHL testosterone (*) aromatase corticosterone cortisol 18hydrxylase estradiol 18-hydroxycorti... glucocorticoid estrogen / androgen 18dehydrogen... mineralocorticoid aldosterone (*) metyperone test androgens CC cp450 ACTH, AgII and LH stimulate CC or desmolase, a rate-limiting enzyme for steroidogenesis teroid and enzymes VLDL cholesterol mevalonic acid AcCoA pyruvate HMG-CoA reductase CC pregnenolone 17 a-hydroxypreg... DHA 17alpha 17alph 3BD hydroxylase 3BDI desmolase a I 3BDI progesterone 17 a-progesterone androstenedione 21BHL 21BHL ßhydroxysteroid dehydrogenase 11-deoxycorti deoxycortisol 11BHL 11BHL testosterone (*) aromatase corticosterone cortisol 18hydrxylase estradiol 18-hydroxycorti... glucocorticoid estrogen / androgen 18dehydrogen... mineralocorticoid aldosterone (*) metyperone test primary (A) and secondary (B, C) regulation sites for steroid biosynthesis Page 3
4 Control of steroid biosynthesis VLDL cholesterol mevalonic acid AcCoA pyruvate HMG-CoA reductase CC pregnenolone 17 a-hydroxypreg... DHA 17alpha 17alph 3BD hydroxylase 3BDI desmolase a I 3BDI progesterone 17 a-progesterone androstenedione 21BHL 21BHL ßhydroxysteroid dehydrogenase 11-deoxycorti deoxycortisol 11BHL 11BHL testosterone (*) aromatase corticosterone cortisol 18hydrxylase estradiol 18-hydroxycorti... glucocorticoid estrogen / androgen 18dehydrogen... mineralocorticoid aldosterone (*) metyperone test sterified (C) and free cholesterol (FC) before, during and after stimulation Control of steroid action gene, exons, introns cis - acting, trans - acting transcription, translation splicing RNA cap polya tail Hormonal action can be regulated at the level of transcription, translation, RNA turnover, protein turnover, and post - translational modification. teroid action can be regulated at each level of its mechanism of action Page 4
5 Mechanism of steroid action NH 2 1 DNA binding region Hormone binding region COOH 563 estrogen progesterone glucocorticoid thyroid hormone androgen 1 teroid action is initiated by the binding of the steroid to its intracellular receptor 427 Vitamin D Mechanism of steroid action teroid action is initiated by the binding of the steroid to its intracellular receptor Page 5
6 Mechanism of steroid action H R HP 90 HP 90 Zn fingers are binding regions of transcription factor proteins which attach to the promotor segment of DNA H R Zn HP 90 HP 90 teroid - receptor complex bind to specific hormone receptive elements on DNA H R Zn H R H R Zn D N A TATA box RNA Pol II teroid s permissive effect R Na / K pump camp ----> PKA ----> channel / enzyme AC Protein synthesis XX1 HR mrna R teroid + R ----> R DNA additional transcription factor teroids, by their genomic effects, can control the availability of specific proteins used by hormones Page 6
7 teroid s permissive effect Action on specific mrna synthesis could cause an increase in the number of membrane receptors, which might increase the production of cyclic nucleotides, thus leading to an increase cellular response to hormones acting on the plasmalemma. Thyroid or steroid hormones could increase or decrease the amount of cyclic nucleotide - dependent protein kinases PK or amount of substrate available for phosphorylation by camp or cgmp - dependent PK. Thyroid and steroid hormones could enhance the synthesis of a protein that could act as an inhibitor of another protein (e.g.. phosphoprotein phosphatase) whose action is antagonistic to cyclic nucleotide action. teroids by their genomic effects can control the availability of specific proteins used by hormones teroid s developmental effect steroid hormones are synthesized and secreted by the ovary, testes, adrenal and placenta their effects occur through binding to intracellular receptors regulating transcription in target cells are critical for sexual differentiation of different target organs and are important regulators (e.g.. they induce developing of Wolffian system, a noncycling hypothalamus and sexual behavior in males, since dogma states default system is female) liver cells express R and AR. They regulate production of secreted proteins (HBG) and enzymes (15ßH) their synthesis / receptors alterations result in disease. teroid-dependent tumors can become independent teroids by their genomic effects can control the development of specific systems and processes Page 7
8 teroid s physiological effect glucocorticoids secreted from adrenal cortex (ZF, cortisol) regulate gluconeogenesis and participate in inflamatory and immune processes mineralocorticoids secreted from adrenal cortex (ZG, aldosterone) is part of the RA system regulating blood pressure and kidney functions gonadal steroids (testis/ovaries) participate in sexual differentiation of reproductive system, its control, and phenotypical gender differences neurosteroids produced in CN and PN (mainly but not exclusively) in myelinating glial cells, from cholesterol or steroidal precursors imported from peripheral sources. ome of them are pregnenolone, progesterone, and DHA. They can act as modulators of GABA and NMDA receptors teroids by their genomic effects can control the function of specific processes teroid s pathological effect Theoretically, genetic pathologies can be associated with each step of the biosynthetic leading to the production of a particular enzyme or protein. congenital adrenal hyperplasia due to gene deletion or to point mutation of the 21 - hydroxylase enzyme testicular feminization due to point mutations scattered throughout the androgen receptor gene, cause decrease amounts of functional androgen receptors, altered sexual differentiation and feed - back regulation Vit D - dependent rickets due to a single point mutation in the tip of one of the Zn fingers of the DNA binding domain of the Vit D receptor, thus making it unable to interact and transcriptionally regulate Vit D - responsive genes teroids by their genomic effects can control the availability of specific proteins used by hormones Page 8
9 Anatomy of the adrenal gland Chemical communication between cells. 1. A: Local. ecretors product, shown as red dots, reaches nearby target cell by diffusion through extracellular fluid (paracrine or autocrine communication). Juxtacrine: Communication by physical contact via signaling molecules in the membrane of one cell activating membrane receptor molecules in an adjacent cell. 2. B: ndocrine. ecretory product reaches distant cells by transport through the circulation. 3. C: ecretory product released from terminals of long cell processes reaches target cells distant from the nerve cell body by diffusion across the synaptic cleft. The adrenal has a cortex made of three distinctly separated regions and a medulla Histology of the adrenal gland The principal adrenal steroid hormones. ZG Aldosterone ZF Cortisol ZR Androgens ach of the three sections of the adrenal cortex, called zonas, produce different hormones Page 9
10 Histology of the adrenal gland CC Conversion of cholesterol to pregnenolone, the rate determining reactions in steroid hormone biosynthesis. Carbons 20 and 22 are sequentially oxidized (in either order) followed by oxidative cleavage of the bond between them (green arrow). All three reactions are catalyzed by a single enzyme, cytochrome P450CC. The principal adrenal steroid hormones. The first reaction in the formation of all adrenal steroids is removal of the side-chain of cholesterol Histology of the adrenal gland Biosynthesis of adrenal cortical hormones. Reactions shown in the yellow box are unique to the zona glomerulosa. Reactions shown in the blue box are seen in the zonae fasciculata. Reactions shown in the green box are seen in both the zona glomerulosa and reticularis. Reactions shown in the pink box Cortisol is the main glucocorticoid in humans, while in are largely confined to the zona reticularis. animals it is cortocosterone tructural changes produced in each reaction are shown in red. ome enzymatic reactions are restricted to different cortical segments, thus producing different steroids Page 10
11 The HPA axis Hypothalamic - pituitary - adrenal axis (HPA axis) is control predominantly by cortisol negative feedback The HPA axis Hypothalamic - pituitary - adrenal axis (HPA axis) is control predominantly by cortisol negative feedback Page 11
12 The zona fasciculata treatment? ZF, glucocorticoids, cortisol, corticosterone stimulus for cortisol secretion is ACTH main effects of cortisol are 1) facilitate gluconeogenesis in muscle and connective tissue; 2) decreases glucose utilization by inhibiting its transport; 3) reduces utilization of aa everywhere except liver; 4) increases mobilization of FA and glycerol from adipose tissue to blood; 5) anti-inflamatory cortisol is needed to mantain vascular integrity / responsiveness and the volume of body fluids. ffect on GFR ZR, androgens, DHA. ACTH rises DHA secretion, but this steroids does not inhibits ACTH secretion from AP in females, a virilization syndrome occurs due to hypersecretion of adrenal androgens caused by a blockade, or deficiency, of the enzyme 21-ß-hydroxylase The zona fasciculata, the second cortical region, secretes the glucocorticoid steroid cortisol The zona fasciculata cortisol dexamethasone treatment? Cortisol is the natural glucocorticoid hormone in humans and dexamethasone is a synthetic analog Page 12
13 Mechanism of action of CRH Hormonal interactions that regulate ACTH secretion by pituitary corticotrope. Glucocorticoids produce their negative feedback effects by interfering with POMC gene expression and membrane depolarization. CRH = corticotrophin releasing hormone; AVP = arginine vasopressin; AC = adenylyl cyclase; PLC = phospholipase C; ATP = adenosine triphosphate; camp = cyclic adenosine monophosphate; PKC = protein kinase C; DAG = diacylglycerol; IP3 = inositol trisphosphate; PKA = protein kinase A; CRB = cyclic AMP response element binding protein; R = endoplasmic reticulum; AP-1 = activator protein-1; GR = glucocorticoid receptor; POMC = proopiomelanocortin; VCC = voltage sensitive calcium channels. The main stimulus for ACTH secretion is hypothalamic CRH (corticotropic releasing hormone) Mechanism of action of ACTH The main stimulus for cortisol secretion is ACTH (adrenocorticotropic hormone) from the AP Page 13
14 Mechanism of action of ACTH timulation of steroidogenesis by ACTH in zona fasciculata cells. Conversion of cholesterol to pregnenolone requires mobilization of cholesterol from its storage droplet and transfer to the P450scc (side chain cleavage) enzyme on the inner mitochondrial membrane. ACTH may also increase cholesterol uptake by increasing the number or affinity of low density lipoprotein (LDL) receptors. s = stimulatory subunit of the guanine nucleotidebinding protein. AC = adenylyl cyclase, = subunits of the guanine nucleotide-binding protein. tar = steroid acute regulatory protein. The main stimulus for cortisol secretion is ACTH (adrenocorticotropic hormone) from the AP Mechanism of action of cortisol The main inhibitor of the HPA axis is the release of cortisol from the adrenal gland Page 14
15 Mechanism of action of cortisol Average plasma concentrations of cortisol and dehydroepiandrosterone sulfate (DHA) throughout life. DHA is abundant in fetal plasma and declines precipitously after birth as the fetal zone of the adrenal involutes. DHA increases during adrenarche as the zona reticularis increases in mass. Although both the ovaries and testes contribute to circulating DHA, the difference in plasma levels between males and females reflects the greater contribution of the testes. Cortisol is secreted by the fetal adrenal cortex only in the latter part of pregnancy. Average adult levels are reached shortly after birth and remain within a constant range throughout life. Cortisol and DHA release to circulation throughout life in humans Basal release of adrenal steroid CORT ALD DHA DHA sulphate The secretion of adrenal steroids to plasma is episodic in nature Page 15
16 Basal release of adrenal steroid The secretion of adrenal steroids to plasma has a circadian rhythm The metyrapone test The metyperone test checks the functionality of the cortisol negative feedback Page 16
17 The metyrapone test The cortisol-cortisone shuttle. Two enzymes, 11-hydroxysteroid dehydrogenase (HD I and HD II) catalyze the oxidation of cortisol to cortisone. HD I can also catalyze the reaction in the reverse direction converting the inactive cortisone to cortisol. Cortisol is the main glucocorticoid in humans while corticosterone is the main one in most animals Insulin induced hypoglycemia The insulin - induced hypoglycemic test is used to verify the responsiveness of the HPA axis Page 17
18 ffects Metabolic effects of cortisol Principal effects of glucocorticoids on glucose production and the metabolism of body fuels. The main metabolic effect of cortisol on metabolism is gluconegenesis Page 18
19 Metabolic effects of cortisol increases gluconeogenesis by a genomic mechanism inhibits glucose transport thus decreases its utilization reduces aa use for protein formation except in the liver increases FA / glycerol mobilization from adipose depot The main metabolic effect of cortisol on metabolism is gluconegenesis Metabolic effects of cortisol glycogen phosphorilase G-6-Pase glucose F-1,6-diPase proteins transaminases amino acids glycogen glycogen synthetase G P hexokinase G P glucose insulin F P glucagon PFK epinephrine F - 1,6 -dip glucocorticoids triose fats pyruvate acetyl CoA increases gluconeogenesis by a genomic mechanism inhibits glucose transport thus decreases its utilization Krebs cycle reduces aa use for protein formation except in the liver increases FA / glycerol mobilization from adipose depot The main metabolic effect of cortisol on metabolism is gluconegenesis Page 19
20 Metabolic effects of cortisol increase synthesis of gluconeogenic enzymes within hepatocytes (anabolic) actions on skeletal muscle and adipose are catabolic (block glucose uptake, stimulate proteolysis, lipolysis, and promote FFA and glycerol mobilization) glucose produced is either stored as glycogen or released into the blood excessive secretion is antagonistic to insulin and promotes diabetes mellitus, an effect that is amplified since they reduce the affinity of certain cells for insulin, further aggravating the diabetes Gluconeogenesis The main metabolic effect of cortisol on metabolism is gluconegenesis Other effects of cortisol most if not all effects are permissive to actions of other hormones. xamples: sympathoadrenal function ADX, stress, vascular collapse CAs, enzymes and fat mobilization lethal hypothermia follows ADX permissive actions mantain body temperature through their permissive actions on liver gluconeogenesis and adipose tissue metabolism sheep fetus and the onset of parturition role of cortisol and Prl in lactogenesis milk casein drops 85% after ADX repro actions Due to its genomic mechanism of action, cortisol has a variety of effects Page 20
21 Other effects of cortisol neonatal corticosterone reduces basal level and amplitude of its diurnal rhythm in adult rat (imprinting, brain diff/dev) hippocampus granular cell layer is loss after ADX and corticosterone prevents it non-genomic mechanisms of steroid action in neuronal function and behavior CN actions at higher than physiological levels, cortisol inhibits inflammatory & allergic reactions, probably by lysosomal membrane stabilization cortisol causes lymphatic atrophy and lymphocytopenia ACTH stimulates IL-1. anti-inflamatory and immune actions Due to its genomic mechanism of action, cortisol has a variety of effects Anti-inflamatory effect Anti-inflammatory actions of cortisol. Cortisol induces the formation of the inhibitor of nuclear factor B (I-B), which binds to nuclear factor B (NF-B) and prevents it from entering the nucleus and activating target genes. The activated glucocorticoid receptor (GR) also interferes with NF-B binding to its response elements in DNA thus preventing the induction of phospholipase A2 (PLA2), cyclooxygenase 2 (COX2), and the inducible nitric oxide synthase (ino). By blocking further production of TNF (tumor necrosis factor-) and IL-1 (interleukin-1) glucocorticoids disrupt the positive feedback cycle involving these cytokines. NO = nitric oxide. Cortisol induces formation of the nuclear factor I-kB which binds NF-kB and prevents its nuclear entry Page 21
22 Anti-inflamatory effect Negative feedback regulation of the HPA axis by inflammatory cytokines Cortisol & the tress Response tress stimulates the HPA axis in both an acute and chronic mode Page 22
23 Cortisol & the tress Response HPA axis recovery from Cushing or chronic DX suggests chronic stress effect Cortisol & the tress Response When threat-related information reaches the brain two different response are generated: 1) The brain automatically engages TH AMIGDALA, an emergency fear center. This sent emergency information to other parts of the brain mediating the stress response (pi, sweat, rapid heart beat, increased blood pressure). 2) The brain engages the THALAMU first and then the CORTX. The latter analyzes the information and decides whether or not a stress response is required. If the threat is not real or the response triggered by the amigdala is excessive, prefrontal cortex inhibits the amigdala. If the threat is real the cortex support it and the body stays on alert. HPA axis recovery from Cushing or chronic DX suggests chronic stress effect Page 23
24 Cortisol & the tress Response The amygdala provokes a series of changes in the brain s chemicals that puts the entire body in a stress mode. The amigdala does not check for accuracy of the threat. Another part of the stress signal goes to the cortex, the thinking part, to figure out what is going on. Then the information goes to the prefrontal cortex, and from here the amygdale receives a very clear message to relax. (This is what supposed to happen in a normal stress response). Cortical assessment of incoming stress signals are compared to those in memory associated with the hypocampus. All rational cortical integration is conveyed to the prefrontal cortex to provide a breaking signal to the emotional outburst from the amygdala. For a normal stress response all circuitry has to be functional HPA axis recovery from Cushing or chronic DX suggests chronic stress effect Cushing yndrome Hyperfunction and hypofunctions of primary or secondary origin Page 24
25 Cushing yndrome Hyperfunction (Cushing): obesity, fat redistribution, hypertension, proximal muscle weakness, loss of connective tissue support with abdominal striae and easy bruisibility, emotional symptoms, osteopenia, glucose intolerance Cushing yndrome is associated with hypersecretion of cortisol from the adrenal Cushing yndrome ACTH Cushing disease Adrenal tumor ctopic ACTH Hyperfunction (Cushing): obesity, fat redistribution, hypertension, proximal muscle weakness, loss of connective tissue support with abdominal striae and easy bruisibility, emotional symptoms, osteopenia, glucose intolerance Cushing yndrome is associated with hypersecretion of cortisol from the adrenal Page 25
26 Addison Disease Hypofunction (Addisson): weight loss, fatigue, weakness, anorexia, postural hypotension. If adrenal gland is absent or destroyed, in addition hyperpigmentation, hypokalemic alkalosis and virilization in women is present A patient with Addison s disease before (note thin and pigmented face and hand) and after treatment with glucocorticoid replacement therapy. Addison Disease is associated with hyposecretion of cortisol from the adrenal Addison Disease ACTH primary adrenal failure secondary adrenal failure Hypofunction (Addisson): weight loss, fatigue, weakness, anorexia, postural hypotension. If adrenal gland is absent or destroyed, in addition hyperpigmentation, hypokalemic alkalosis and virilization in women is present Addison Disease is associated with hyposecretion of cortisol from the adrenal Page 26
27 Addison Disease Page 27
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