Chapter 26 The Urinary System The Urinary System Copyright © John Wiley & Sons, Inc. All rights reserved. Renal Anatomy § The kidneys are located between the peritoneum and the posterior wall of the abdomen (in the retroperitoneal space). l They are partially protected by the eleventh and twelfth pairs of ribs. l Because of the position of the liver, the right kidney is slightly lower than the left. Copyright © John Wiley & Sons, Inc. All rights reserved. Transverse plane ANTERIOR Large intestine Stomach Pancreas Liver Renal artery and vein View Layers Inferior vena cava Peritoneum Abdominal aorta RENAL HILUM Body of L2 RENAL FASCIA LEFT KIDNEY ADIPOSE CAPSULE RENAL CAPSULE Spleen Rib RIGHT KIDNEY POSTERIOR (a) Inferior view of transverse section of abdomen (L2) Quadratus lumborum muscle Location & layers Copyright © John Wiley & Sons, Inc. All rights reserved. Renal Anatomy § A frontal section through the kidney reveals two distinct regions of internal anatomy, the cortex and medulla. l The main function of the cortex is filtration to form urine. l The main function of the medulla is to collect and excrete urine. Copyright © John Wiley & Sons, Inc. All rights reserved. PATH OF URINE DRAINAGE: Nephron Collecting duct Renal hilum Minor calyx Renal cortex Renal medulla Renal artery Renal vein Major calyx Renal pelvis Renal column Renal pyramid in renal medulla Renal papilla Ureter Renal capsule Urinary bladder (a) Anterior view of dissection of right kidney Copyright © John Wiley & Sons, Inc. All rights reserved. Renal Anatomy § The renal pyramids within the medulla contain the kidney’s tubules. The renal papilla is the location where the pyramids empty urine into cuplike structures called minor calyces. 2-3 minor calyces empty into a major calyx § From the major calyces, urine drains into § the renal pelvis and then out through the ureter Copyright © John Wiley & Sons, Inc. All rights reserved. Renal Blood Flow § The renal artery and renal vein pass into the substance of the kidney at the hilum. ◦ The renal arteries are very large branches of the aorta, and up to a third of total cardiac output can pass through them to be filtered by the kidneys. Copyright © John Wiley & Sons, Inc. All rights reserved. Copyright © John Wiley & Sons, Inc. All rights reserved. Renal Blood Flow Copyright © John Wiley & Sons, Inc. All rights reserved. The Nephron § Nephrons are the structural and functional units that form urine 1 million in each kidney- consist of: l Renal corpuscle associated with a l Renal tubule Copyright © John Wiley & Sons, Inc. All rights reserved. The Nephron: Renal Corpuscle § The Renal Corpuscle consists of two structures: l The glomerular capillaries l The Bowman’s capsule– a double-walled epithelial cup that surrounds the glomerular capillaries. Copyright © John Wiley & Sons, Inc. All rights reserved. § Each nephron receives one afferent arteriole, which divides into a tangled, ball-shaped capillary network called the glomerulus. l The glomerular capillaries then reunite to form an efferent arteriole that carries blood out of the glomerulus. Copyright © John Wiley & Sons, Inc. All rights reserved. § Bowman’s capsule consists of visceral and parietal layers. l The visceral layer is made of epithelial cells called podocytes. l The parietal layer of the glomerular capsule is a simple squamous epithelium and forms the outer wall of the capsule. l Fluid filtered from the glomerular capillaries enters Bowman’s space, (the space between the two layers of the glomerular capsule) Copyright © John Wiley & Sons, Inc. All rights reserved. Renal corpuscle (external view) Parietal layer of glomerular (Bowman’s) capsule Afferent arteriole Mesangial cell Juxtaglomerular cell Capsular space Macula densa Ascending limb of loop of Henle Proximal convoluted tubule Mesangial cell Efferent arteriole Podocyte of visceral layer of glomerular (Bowman’s) capsule Endothelium of glomerulus Pedicel (a) Renal corpuscle (internal view) Copyright © John Wiley & Sons, Inc. All rights reserved. Glomerular capsule: Glomerulus Parietal layer Podocytes of visceral layer of glomerular capsule Visceral layer Afferent arteriole Juxtaglomerular cell Capsular space Ascending limb of loop of Henle Macula densa cell Simple squamous epithelial cells Efferent arteriole Proximal convoluted tubule LM 1380x (b) Renal corpuscle Copyright © John Wiley & Sons, Inc. All rights reserved. The Nephron: Tubule l Filtered fluid passes into the renal tubule, which has three main sections: ◦1. the proximal convoluted tubule (PCT) ◦2.the loop of Henle; consists of: ◦the descending limb ◦the ascending limb ◦3.the distal convoluted tubule (DCT) Copyright © John Wiley & Sons, Inc. All rights reserved. Microvilli Lining epithelium Mitochondrion Apical surface Proximal convoluted tubule (PCT) Loop of Henle: descending limb and thin ascending limb Loop of Henle: thick ascending limb Most of distal convoluted tubule (DCT) Last part of DCT and all of collecting duct (CD) Intercalated cell Principal cell Copyright © John Wiley & Sons, Inc. All rights reserved. The Nephron § The distal convoluted tubules of several nephrons empty into a single collecting duct. l Collecting ducts unite and converge into large papillary ducts which drain into the minor calyces. Copyright © John Wiley & Sons, Inc. All rights reserved. The Nephron § Nephrons can be sorted into two populations: cortical nephrons ( 85%) with short loops of Henle their blood supply is from peritubular capillaries that arise from efferent arterioles. juxtamedullary nephrons ( 15%): § nephrons with long loops of Henle enable the kidneys to create a concentration gradient in the renal medulla and to excrete concentrated urine. Their loops of Henle receive their blood supply from the vasa recta Copyright © John Wiley & Sons, Inc. All rights reserved. The Cortical Nephron Copyright © John Wiley & Sons, Inc. All rights reserved. Juxtamedullary Nephron Copyright © John Wiley & Sons, Inc. All rights reserved. Juxtaglomerular Apparatus (JGA) § JGA is located where the distal tubule lies near the afferent arteriole § JGA consists of: § 1.Juxtaglomerular (JG) cells)- in the wall of afferent arterioles: l l l Are enlarged, smooth muscle cells Have secretory granules containing renin Sensitive to blood pressure changes § 2.Macula densa l l l Tall, distal tubule cells Lie adjacent to JG cells Sensitive to NaCl concentration in filtrate Copyright © John Wiley & Sons, Inc. All rights reserved. Juxtaglomerular Apparatus (JGA) § The JGA helps regulate blood pressure within the kidneys. Copyright © John Wiley & Sons, Inc. All rights reserved. Kidney functions § Removes toxins, nitrogenous metabolic wastes ( urea, uric acid, creatinine) § Regulates blood volume & blood pressure (produces Renin) § Maintains electrolyte balance § Maintains acid base balance § Role in erythropoiesis (produces EPO) Copyright © John Wiley & Sons, Inc. All rights reserved. Mechanisms of Urine Formation § Urine formation involves three major processes: l l l 1.Glomerular filtration 2.Tubular reabsorption 3.Secretion Copyright © John Wiley & Sons, Inc. All rights reserved. Glomerular Filtration: Filtration Membrane § Composed of three layers l Fenestrated endothelium of the glomerular capillaries- blocks formed elements l Basement membrane between the two layers- blocks large proteins l Podocytes slit membranes l (visceral layer of the glomerular capsule)blocks intermediate sized proteins Copyright © John Wiley & Sons, Inc. All rights reserved. Glomerular Filtration § Glomerular filtration is the formation of a protein-free filtrate (ultrafiltrate) of plasma across the glomerular membrane. l Only a portion of the blood plasma delivered to the kidney via the renal artery is filtered. l Plasma which escapes filtration, along with its protein and cellular elements, exits the renal corpuscle via the efferent arterioles l Copyright © John Wiley & Sons, Inc. All rights reserved. Glomerular Filtration A passive process, by which fluids and solutes are pushed through the filtration membrane § The kidneys produce 180L of filtrate/day § The glomerulus is more efficient filter than other capillary beds because: l Its filtration membrane has higher surface area & is more permeable l Glomerular blood pressure is higher- higher net filtration pressure § Blood cells & plasma proteins not filtered (occasional small proteins may leak out) Copyright © John Wiley & Sons, Inc. All rights reserved. Glomerular Filtration § Filtration is controlled by Starling forces. l Blood hydrostatic pressure (55mmHg) is the main force that “pushes” water and solutes through the filtration membrane - (this is the blood pressure in glomerular capillaries) l Capsular hydrostatic pressure (15 mmHg) is exerted against the filtration membrane by fluid in the capsular space (opposes filtration). l Colloidal osmotic pressure (30 mmHg) is the pressure of plasma proteins “pulling” on water (opposes filtration). Copyright © John Wiley & Sons, Inc. All rights reserved. Net Filtration Pressure Net Filtration pressure = Blood Hydrostatic Pressure – Blood Osmotic Pressure – Capsular Hydrostatic Pressure Copyright © John Wiley & Sons, Inc. All rights reserved. Net Filtration pressure = 55-30-15 = 10 mmHg Glomerular Filtration Rate (GFR) The total volume of filtrate formed per minute by the kidneys- 120-125ml/min GFR mainly determined by: •Net filtration pressure- any change in the 3 pressures influences NFP & therefore GFR •Changes in GFR normally result from changes in blood hydrostatic pressure (glomerular blood pressure) Copyright © John Wiley & Sons, Inc. All rights reserved. Regulation of GFR § Regulation of the GFR is critical to maintaining homeostasis and is regulated by local and systemic mechanisms: l Renal autoregulation occurs when the kidneys themselves regulate GFR. l Neural regulation occurs when the sympathetic nerve supply regulates renal blood flow and GFR. l Hormonal regulation involves angiotensin II and atrial natriuretic peptide (ANP). Copyright © John Wiley & Sons, Inc. All rights reserved. Renal Autoregulation of GFR § Under normal conditions, renal autoregulation maintains a constant GFR despite fluctuations in BP § Two local mechanisms in the kidney that control GFR (Renal autoregulation) ◦ Myogenic mechanism ◦ Tubuloglomerular feedback § Myogenic – VC of afferent arteriole in response to increase in BP, & VD in response to fall in BP, therefore maintaining GFR Copyright © John Wiley & Sons, Inc. All rights reserved. Renal Autoregulation of GFR § Tubuloglomerular feedback – macula densa cells of JGA respond to NaCl concentration in the filtrate § As GFR increases- less time for Na reabsorption- high NaCl concentration in filtrate § macula densa cells release VCs--- VC of afferent arteriole ----leading to fall in GFR Copyright © John Wiley & Sons, Inc. All rights reserved. Tubuloglomerular feedback Copyright © John Wiley & Sons, Inc. All rights reserved. Regulation of GFR § Neural regulation of GFR is possible because the renal blood vessels are supplied by sympathetic fibers that release norepinephrine causing vasoconstriction. § Decreases GFR l Sympathetic input to the kidneys is most important with extreme drops of B.P. (as occurs with hemorrhage). Copyright © John Wiley & Sons, Inc. All rights reserved. Regulation of GFR § Two hormones contribute to regulation of GFR l Angiotensin II is a potent vasoconstrictor of both afferent and efferent arterioles (reduces GFR). l Atrial natriuretic peptide (ANP). l A sudden large increase in BP stretches the cardiac atria and releases ANP causes the afferent arterioles to dilate & glomerulus to relax, increasing the surface area for filtration. (increases GFR). Copyright © John Wiley & Sons, Inc. All rights reserved. Tubular Reabsorption § Tubular reabsorption is the process of returning important substances (“good stuff”) from the filtrate back into the renal blood vessels... and ultimately back into the body. Copyright © John Wiley & Sons, Inc. All rights reserved. Tubular Reabsorption § The “good stuff” is glucose, electrolytes, vitamins, water, amino acids, and any small proteins that might have escaped from the blood into the filtrate. § Ninety nine percent of the glomerular filtrate is reabsorbed (most of it before the end of the PCT)! Copyright © John Wiley & Sons, Inc. All rights reserved. Tubular Reabsorption Amount in 180 L of filtrate (/day) Amount returned to blood/d (Reabsorbed) Amount in Urine (/day) 3L 180 L 178-179 L 1-2 L Protein (active) 200 g 2g 1.9 g 0.1 g Glucose (active) 3g 162 g 162 g 0g Urea (passive) 1g 54 g 24 g 30 g (about 1/2) (about 1/2) 0.03 g 1.6 g 0g 1.6 g (all filtered) (none reabsorbed) Total Amount in Plasma Water (passive) Creatinine Copyright © John Wiley & Sons, Inc. All rights reserved. Tubular Reabsorption § Reabsorption into the interstitium has two routes: l Paracellular reabsorption is a passive process that occurs between adjacent tubule Cells Transcellular reabsorption is movement through an individual cell.( must pass through apical & basal membranes) Copyright © John Wiley & Sons, Inc. All rights reserved. Tubular Reabsorption; transport mechanisms Reabsorption of fluids, ions, and other substances occurs by active and passive means including: Diffusion Osmosis (water)- from high water to low water ( from low osmolarity to high osmolarity) Primary active transport- energy derived from ATP is used to “pump” a substance across a membrane Secondary active transport- energy stored in an ion’s electrochemical gradient drives another substance across the membrane. Copyright © John Wiley & Sons, Inc. All rights reserved. Transport mechanisms Reabsorption of water can be obligatory or facultative Obligatory reabsorption of water occurs when it is obliged to follow solutes as they are reabsorbed Occurs in PCT( always permeable to water) Facultative reabsorption (upto 10% of total water reabsorption) is variable water reabsorption, adapted to specific needs. It is regulated by of ADH on the principal cells of DCT & collecting ducts (without ADH they are not permeable to water) Copyright © John Wiley & Sons, Inc. All rights reserved. Reabsorption in the Proximal Convoluted Tubule The majority of solute and water reabsorption from filtered fluid occurs in the PCT which reabsorbs, 65% Na+, water, K+, 80-90% of bicarbonate ions Most absorptive processes involve Na+reabsorption PCT Na+ reabsorption promotes reabsorption of; water, K+; Cl- and other ions Normally, 100% of filtered glucose, amino acids, and other nutrients are reabsorbed in the PCT by Na+ symporters. ( co-transported with sodium) Copyright © John Wiley & Sons, Inc. All rights reserved. Reabsorption of glucose due to active pumping of Na+ Na+ symporters help reabsorb materials from the tubular filtrate Glucose is completely reabsorbed in the first half of the PCT Sodium levels are kept low inside the cells due to Na+/K+ pump---- Reabsorption of Nutrients sodium moves in from tubular fluid down the gradient, cotransporting glucose Copyright © John Wiley & Sons, Inc. All rights reserved. Transport Maximum A transport maximum (Tm) is maximum reabsorption rate, i.e. rate when all the carriers are saturated- in mg/min When the carriers are saturated, excess of that substance is excreted in urine In DM, due to hyperglycemia-when all carriers are saturated glucose starts appearing in urine--glucosuria Copyright © John Wiley & Sons, Inc. All rights reserved. Reabsorption of water, other ions, due to active pumping of Na+ Water follows sodium by osmosis through aquaporins in PCT (obligatory water reabsorption) Passive diffusion of anions Cl-, K+, Ca++, to maintain electroneutrality Urea, fat-soluble substances- move down their gradient as filtrate becomes more concentrated due to PCT water movement Copyright © John Wiley & Sons, Inc. All rights reserved. PCT-Reabsorption of Bicarbonate, Na+ & H+ secretion Na+/H+ antiporters reabsorb Na+ and secrete H+ PCT cells produce the H+ & release bicarbonate ion to the peritubular capillaries important buffering system For every H+ secreted into the tubular fluid, one filtered bicarbonate eventually returns to the blood Copyright © John Wiley & Sons, Inc. All rights reserved. Reabsorption in the Loop of Henle Water reabsorption: about 15% of the filtered water is reabsorbed in the descending limb, little or no water is reabsorbed in the ascending limb Solute reabsorption occurs in ascending limb Na+-K+-Cl- symporters reclaim Na+, Cl-, and K+ ions from the tubular lumen fluid in the thick part of ascending limb Copyright © John Wiley & Sons, Inc. All rights reserved. Reabsorption in the Loop of Henle Fluid in tubule lumen Vasa recta Thick ascending limb cell Na+ ATP Main result is reabsorption of Na+ and Cl-. Na+ ADP Na+ 2Cl– K+ Cations: Na+ K+ Ca2+ Mg2+ Na+ Na+ 2Cl– 2Cl– 2Cl– K+ Cations Key: Na+–K+–2Cl– symporter Apical membrane (impermeable to water) Interstitial fluid is more negative than fluid in tubule lumen Leakage channels Sodium–potassium pump Diffusion Copyright © John Wiley & Sons, Inc. All rights reserved. Reabsorption and Secretion in the late DCT & Collecting Ducts DCT & Collecting ducts- most reabsorption depends on needs of the body & regulated by hormones: Aldosterone for Na+ Aldosterone for water (by ADH release ) ADH for water- otherwise impermeable to water PTH for Ca2+ ANP-Acts on CDs to inhibit Na+ reabsorption reduces blood Na+ & BP Copyright © John Wiley & Sons, Inc. All rights reserved. Tubular Secretion Substances move from peritubular capillaries into the filtrate Main site of secretion is PCT (except for K+ ) It is by active transport. Tubular secretion is important for: Disposing of drugs, metabolites- not filtered because bound to plasma proteins Eliminating undesirable substances such as urea, uric acid, creatinine Ridding the body of excess K+ ions ( by aldosterone in DCTs & CDs) + Controlling blood pH- (secreting H+), NH ions 4 © John Copyright Wiley & Sons, Inc. All rights reserved. Hormonal Regulation of Tubular Reabsorption & Secretion Antidiuretic Hormone (ADH) : affects facultative water reabsorption by increasing the water permeability of principal cells in the last part of the distal convoluted tubule and throughout the collecting duct. In the absence of ADH, these are almost impermeable to water. Copyright © John Wiley & Sons, Inc. All rights reserved. Hormonal Regulation Renin- Angiotensin and aldosterone system: When blood volume and blood pressure decreases, JGA secretes renin. Promotes aldosterone production which causes principal cells: ◦ to reabsorb more Na+ followed by water ( through ADH release) ◦ To secrete K+ ions ◦ Decreases urine output & increases blood volume by increasing water reabsorption Copyright © John Wiley & Sons, Inc. All rights reserved. Hormonal Regulation Atrial natriuretic peptide ◦ inhibits reabsorption of Na+ and water ◦ increase excretion of Na+ which increases urine output and decreases blood volume ◦ Parathyroid hormone: ◦ In response to low blood calcium, PTH stimulates cells in DCT to reabsorb more calcium. Copyright © John Wiley & Sons, Inc. All rights reserved. Production of Dilute and Concentrated Urine The rate at which water is lost from the body depends mainly on ADH When ADH level is very low, the kidneys produce dilute urine and excrete excess water When ADH level is high, the kidneys secrete concentrated urine and conserve water; a large volume of water is reabsorbed from the tubular fluid and the solute concentration of urine is high. Copyright © John Wiley & Sons, Inc. All rights reserved. Production of Dilute and Concentrated Urine Production of concentrated urine involves: 1.The presence of ADH which makes the distal part of nephron permeable to water 2.The countercurrent mechanism which creates high osmotic gradient for maximal water reabsorption Copyright © John Wiley & Sons, Inc. All rights reserved. The Countercurrent Mechanism Creates a high osmotic gradient from the cortex to the medulla It includes: Countercurrent multiplication Involves long loops of Henle of juxta medullary nephrons Countercurrent exchange. Involves blood flow in the Vasa recta around the loops of Henle Copyright © John Wiley & Sons, Inc. All rights reserved. The Countercurrent Mechanism Countercurrent multiplication is the process by which a progressively increasing osmotic gradient is formed in the interstitial fluid of renal medulla as a result of countercurrent flow (opposing fluid flow in adjacent limbs of long loops of Henle of juxtamedulary nephrons) Countercurrent exchange is the process by which solutes and water are passively exchanged between the blood of the vasa recta and interstitial fluid of the renal medulla Copyright © John Wiley & Sons, Inc. All rights reserved. The Countercurrent Mechanism The gradient from cortex to medulla is 300-1200mOsm The solute concentration in the PCTs is 300 mOsm Water only leaves descending limbfiltrate osmolarity increases to 1200 Salt only reabsorbed in ascending limb- osmolarity of filtrate decreases to 100 at top of loop Copyright © John Wiley & Sons, Inc. All rights reserved. The Countercurrent Mechanism The descending loop of Henle: Is impermeable to solutes & permeable to water Water passes out by osmosis and tubular fluid increases in osmolarity reaching an equilibrium with the Interstitial fluid of medulla The ascending loop of Henle: Is permeable to solutes & impermeable to water Na Cl is pumped out - tubular fluid becomes progressively more dilute - by the time it reaches the DCT osmolarity is 100 mOsm/L The vasa recta maintain the high osmolarity in the medulla Blood flows down the descending limb then up the ascending limb of Vasa recta- exchanges a lot of fluid and solute- but does not carry away any excess salt Copyright © John Wiley & Sons, Inc. All rights reserved. The vasa recta help maintain the high osmolarity in the medulla; exchanges a lot of fluid and solutebut do not carry away any excess salt H2O H2O H2O H2O Copyright © John Wiley & Sons, Inc. All rights reserved. The Countercurrent Mechanism Contributes to high osmolarity of medulla Copyright © John Wiley & Sons, Inc. All rights reserved. Formation of Dilute Urine Filtrate is dilute at the end of the ascending loop of Henle Dilute urine is created by allowing this filtrate to continue into the renal pelvis This will happen as long as (ADH) is not being secreted Collecting ducts remain impermeable to water; no further water reabsorption occurs Urine osmolarity can be as low as 65 mOsm (one-sixth that of plasma) Copyright © John Wiley & Sons, Inc. All rights reserved. Formation of Concentrated Urine ADH-acts on CDs to cause further reabsorption of water In the presence of ADH, most of the water in filtrate can be reabsorbed High medullary osmolarity provides an osmotic gradient necessary for water reabsorption to occur in the presence of high levels of ADH Small volume of concentrated urine formed (can be 1200mOsm) Copyright © John Wiley & Sons, Inc. All rights reserved. Vasa recta Loop of Henle Juxtamedullary nephron and its blood supply together Glomerular (Bowman’s) capsule H2O Na+CI– Blood flow Glomerulus Afferent arteriole Distal convoluted tubule Presense of Na+-K+-2CI– symporters Interstitial fluid in renal cortex 200 HO H2O 2 Efferent arteriole 300 300 Collecting duct 300 300 100 H2O 320 Na+CI– 400 Interstitial fluid in renal medulla 380 200 H2O 400 3 Principal cells in Osmotic gradient H2O collecting duct reabsorb more water when ADH is present Na+CI– 400 500 H2O 600 H2O 580 600 320 300 H2O Proximal convoluted tubule Flow of tubular fluid 400 H2O Na+CI– 600 1 Symporters in thick ascending limb cause buildup of Na+ and Cl– 800 700 780 600 Urea H2O 980 1000 H2O 800 800 H2O 800 900 4 Urea recycling 1000 causes buildup of urea in the renal medulla Na+CI– H2O 1000 1100 H2O 1200 2 Countercurrent flow through loop of Henle establishes an osmotic gradient 1200 Loop of Henle 1200 Papillary duct 1200 Concentrated urine (a) Reabsorption of Na+CI– and water in a long-loop juxtamedullary nephron 1200 Copyright ©of John Wiley & Sons, All rights (b) Recycling salts and urea in theInc. vasa recta reserved. Copyright © John Wiley & Sons, Inc. All rights reserved. Renal function tests Two blood tests are commonly done clinically to assess the adequacy of renal function. Blood urea nitrogen (BUN) Serum Creatinine levels Renal Clearance: the volume of plasma that is cleared of a particular substance in a given time-used to determine GFR Creatinine clearance test: for approximate measure of GFR Because creatinine is freely filtered and not reabsorbed at all (some more added by secretion) Copyright © John Wiley & Sons, Inc. All rights reserved. Urinary bladder The ureters transport urine from the renal pelvis of the kidneys to the bladder. The urinary bladder is a hollow, distensible muscular organ with a capacity that averages 700800mL. The urethra is a small tube leading from the internal urethral orifice in the bladder floor to the exterior. Copyright © John Wiley & Sons, Inc. All rights reserved. Copyright © John Wiley & Sons, Inc. All rights reserved. Copyright © John Wiley & Sons, Inc. All rights reserved. Micturition Reflex When volume of urine in bladder is between 200 to 400 mLstretch receptors activated --impulses relayed along sensory axons to stimulate micturition reflex center in sacral spinal cord Parasympathetic stimulation causes detrusor muscle to contract and internal urethral sphincter to open • Conscious control of urination • If desired, voluntary signal from the cortex sent – Signals go through pudendal nerve--cause relaxation of external urethral sphincter Copyright © John Wiley & Sons, Inc. All rights reserved.