Web Tutorial 42.

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The Mammalian Kidney
Textbook sections
42.4 Water and Electrolyte Balance in Terrestrial Vertebrates The Structure of the Kidney Filtration: The Renal Corpuscle Reabsorption: The Proximal Tubule Creating an Osmotic Gradient: The Loop of Henle Regulating Water and Electrolyte Balance: The Distal Tubule and the Collecting Duct Describe the basic anatomy of the mammalian kidney. Identify the parts of the nephron.

After reading the text material, you should be able to

After completing this tutorial, you should be able to

Describe the basic process of urine formation in mammals. Relate the processes involved in urine production to specific regions of the nephron.

NARRATION
Overview of the Kidney
Kidneys are bean-shaped, paired organs that regulate water and electrolyte balance in terrestrial vertebrates. They also remove nitrogenous wastes, usually in the form of urea, from the blood and eliminate them in the urine.

The ureter is a long tube that transports the urine formed in the kidney to the bladder for storage prior to excretion.

The renal artery and renal vein are large blood vessels that carry blood to and from each kidney.

Parts of the Nephron

The outer region of the kidney is called the cortex, and the inner region is called the medulla.

The nephron is the basic functional unit of the kidney. Nephrons perform the work required to maintain water and electrolyte balance. Most nephrons are located in the cortex, but some extend into the medulla.

Each nephron is made up of five regions, each performing a different function: 1. 2. 3. The proximal tubule actively reabsorbs electrolytes, nutrients, and water. Twothirds of the electrolytes and water filtered from the blood is returned to the blood through the proximal tubule. The loop of Henle creates a steep osmotic gradient by reabsorbing water and electrolytes differentially along its U-shaped length. This gradient allows the kidney to concentrate urine. A network of blood vessels supplies the loop of Henle. The vessels return water and electrolytes to the blood without disrupting the osmotic gradient. The distal tubule, like some other regions of the nephron, reabsorbs electrolytes and water, but its activity depends on the bodys state of hydration and is under hormonal control. Urine formation begins in the renal corpuscle, which filters the blood.

4. 5.

Urine formation ends in the collecting duct, where more water may be reabsorbed. The collecting duct is also under hormonal control.

The Nephron and Urine Formation

Well use this simplified and flattened image of a nephron to describe the functions of each of its five regions.

The first stage of urine formation begins in the renal corpuscle with the filtration of blood. Blood entering the renal corpuscle flows into a cluster of capillaries called the glomerulus. The structure around the glomerulus is called Bowmans capsule. The glomerular capillaries have pores and are surrounded by cells with slits. These features allow molecules and ions to be filtered from the blood based on their size. The high pressure inside the capillaries pushes water and small solutes out of the pores and slits and into the surrounding capsule, where the pressure is lower. Large components, such as proteins and cells, cannot fit through the pores and slits and remain in the blood, never entering the nephron. The resulting filtrate exits the renal corpuscle. The filtrate formed in the renal corpuscle contains waste products as well as valuable nutrients, electrolytes, and water. This filtrate enters the proximal tubule, where many of the valuable substances are reabsorbed. The epithelium of the proximal tubule has microvilli, which increase the surface area available for reabsorption.

Intracellular Na+ is actively transported out of the cells lining the tubule, a process that uses ATP. This pumping creates an Na+ gradient favoring the entry of Na+ into the cell from the lumen. Na+ moves through cotransporters, which use the Na+ gradient to cotransport nutrientssuch as glucose, vitamins, and chloride ionsfrom the proximal tubule into the epithelial cell. The movement of Na+ down its concentration gradient provides the energy to move the other solutes against their concentration gradients. Water follows by osmosis.

The reabsorbed electrolytes, nutrients, and water enter blood vessels near the tubule and are returned to the body.

Two-thirds of the water and NaCl filtered by the renal corpuscle is returned to the body via the proximal tubule. The osmolarity of the remaining filtrate in the proximal tubule is unchanged, however, because water reabsorption is proportional to solute reabsorption. The filtrate that emerges from the proximal tubule is relatively high in wastes and low in nutrients, and it is greatly reduced in volume. This fluid enters the loop of Henle. The loop of Henle sets up a steep osmotic gradient in itself and in the surrounding tissue. The osmolarity of the filtrate inside the loop is low in the cortex and high in the medulla, and this gradient is mirrored in the surrounding tissue. The loop of Henle functions as a countercurrent exchanger. The term countercurrent refers to the flow of filtrate in opposite directions in the two limbs of the loop of Henle. The osmotic gradient would disappear if the filtrates in these adjacent tubes ran the same waythere could be no exchange of water and solutes with the surrounding tissue.

After the fluid rounds the bend, it enters the thin ascending limb, which is highly permeable to Na+ and Cl and moderately permeable to urea. These solutes exit passively, following their concentration gradients. This part of the loop is impermeable to water, however. In the thick ascending limb, Na+ and Cl ions are actively pumped out of the tubule. Therefore, as the fluid moves up the tubule, it again becomes less concentrated.

Water and solutes move through the loop of Henle in the following way. The descending limb of the loop is highly permeable to water but impermeable to solutes. Water is lost by osmosis as it moves from a region of lower solute concentration inside the tubule to a region of higher solute concentration in the surrounding tissue. The fluid inside the tubule therefore becomes more concentrated as it flows deeper into the medulla.

The loop of Henle is an example of a positive feedback system, in which the product of a process causes the process to accelerate. The positive feedback works like this: 1. 2. 3. The thick ascending limb pumps solutes into the surrounding tissue.

The high concentration of solutes in the surrounding tissue causes water to exit the descending limb by osmosis, leaving a hypertonic fluid inside the tubule.

What happens to the water and salt that move out of the loop? They quickly diffuse into a blood vessel that runs along the loop and returns the water and salt to the rest of the body. This return is especially important for the water that diffuses out of the descending loop, which would otherwise dilute the solutes in the kidney. The processes that we have discussed so farfiltration, reabsorption, and the establishment of an osmotic gradientare, for the most part, not subject to hormonal regulation. The major solutes in the filtrate that exits the loop of Henle are urea and other wastes, and this fluid is fairly constant in composition over time.

This hypertonic fluid then loses solutes passively to the surrounding tissue, thus completing the positive feedback loop.

The filtrate then enters the distal tubule, where electrolytes and water are reabsorbed. Hormones control the amount of electrolytes and water reabsorbed by this part of the kidney. If Na+ levels in the blood are low, the hormone aldosterone is released. Aldosterone stimulates the reabsorption of Na+ and Cl in the distal tubule. Water follows these ions out of the distal tubule via osmosis.

Hormones also control the concentration of the resulting urine within the collecting duct.

Under conditions of dehydration, antidiuretic hormone (ADH) is released. ADH stimulates both the distal tubule and collecting duct to become highly permeable to water. Water is reabsorbed into the surrounding tissue rather than being lost as urine, thus decreasing the volume of water in the urine. Individuals with diabetes insipidus produce little or no ADH. In the absence of ADH, the distal tubule and collecting duct are relatively impermeable to water. The lack of ADH results in the formation of a copious amount of urine that is low in solutes. As a unit, the nephron is remarkably effective in regulating water and electrolyte balance and maintaining homeostasis.

aldosterone A steroid hormone secreted by the adrenal cortex that regulates the salt and water balance of the body. antidiuretic hormone (ADH) A polypeptide hormone secreted by the pituitary gland that increases blood pressure and decreases urine flow. bladder The organ in terrestrial vertebrates where urine is stored before being transported to the body surface by the urethra, then excreted. Bowmans capsule The region of the nephron that surrounds the glomerulus.

KEY TERMS & CONCEPTS

collecting duct A structure of the vertebrate nephron where water may be reabsorbed and urine formation ends. It is under hormonal control. cotransporter A membrane protein that uses the energy released by the entry of a proton along its electrochemical gradient to transport nitrate, phosphate ions, or other anions against their electrochemical gradient. An array of cations also enters cells via cotransporters. cortex The outer region of the terrestrial vertebrate kidney.

countercurrent exchanger A system in which substances passively diffuse across a membrane separating two countercurrent streams, so that at each end, the fluid leaving along one side of the membrane nearly resembles, in composition, the fluid entering the other side. diabetes insipidus A disorder caused by a lack of antidiuretic hormone and characterized by intense thirst and by the excretion of large amounts of urine.

distal tubule The convoluted portion of the vertebrate nephron that lies between the loop of Henle and the nonsecretory part of the nephron; its function is to reabsorb electrolytes and water and to concentrate urine. It is under hormonal control. electrolyte A compound that dissociates into ions when dissolved in water. epithelium A class of animal tissues consisting of layers of tightly packed cells that line an organ, duct, or body surface. filtrate A pre-urine resulting from the filtration of blood by the glomerulus. glomerulus A cluster of or capillaries that bring blood to the nephron from the renal artery. homeostasis The relatively constant chemical and physical conditions in the cells, tissues, and organs of an animal. kidney The organ that regulates water and electrolyte balance in terrestrial vertebrates. loop of Henle A U-shaped part of a vertebrate nephron that lies between, and is continuous with, the proximal and distal tubules. The loop of Henle creates a steep osmotic gradient by reabsorbing water and electrolytes differentially. lumen The cavity inside any tube or sac-shaped organ. medulla The inner region of the terrestrial vertebrate kidney.

nephron One of the tiny tubes within the vertebrate kidney that filters blood and concentrates salts to produce urine.

microvillus (plural: microvilli) A tiny protrusion from the surface of an epithelial cell that increases the surface area for absorption of substances. Microvilli are common in certain animal organs, such as the vertebrate kidney and intestine. osmolarity The concentration of dissolved substances in a solution, measured in moles per liter.

positive feedback The stimulation of a process as a result of an end product of the process. For example, the presence of an osmotic gradient stimulates water and ion flows that in turn create an osmotic gradient. proximal tubule The convoluted portion of the vertebrate nephron that lies between the renal corpuscle and the loop of Henle and reabsorbs electrolytes, nutrients, and water from the glomerular filtrate.

osmosis The diffusion of water across a selectively permeable membrane from a region of high water concentration (or low solute concentration) to a region of low water concentration (or high solute concentration).

renal corpuscle A structure in the nephron, consisting of the glomerulus and Bowman's capsule, that filters the blood to remove nitrogenous wastes in terrestrial vertebrates. urea A soluble, weakly basic nitrogenous compound that is the chief solid component of mammalian urine and an end product of protein decomposition. In humans, enzyme-catalyzed reactions convert ammonia to urea, which is a much less toxic compound than ammonia.

ureter A long tube in terrestrial vertebrates that transports urine from the kidney to the bladder.