28 Chapter 28: Osmoregulation and Urinary Systems

Joshua Reid

Learning Objectives

By the end of this chapter, you should be able to:

28.1 Define osmosis and the role it plays in maintaining homeostasis in different organisms

28.2 Explain why osmoregulation and osmotic balance are important body functions

28.3 Describe osmoregulators or osmoconformers and how these organisms have adapted to different environments

28.4 Explain how the kidneys serve as the main osmoregulatory organs in the human body

The daily intake recommendation for human water consumption is eight to ten glasses of water. In order to achieve a healthy balance, the human body should excrete the eight to ten glasses of water every day. This occurs via the processes of urination, defecation, sweating and, to a small extent, respiration. The organs and tissues of the human body are soaked in fluids that are maintained at constant temperature, pH, and solute concentration, all crucial elements of homeostasis. The solutes in body fluids are mainly mineral salts and sugars, and osmotic regulation is the process by which the mineral salts and water are kept in balance. Osmotic homeostasis is maintained despite the influence of external factors like temperature, diet, and weather conditions.

Need for Osmoregulation

Biological systems constantly interact and exchange water and nutrients with the environment by way of consumption of food and water and through excretion in the form of sweat, urine, and feces. Without a mechanism to regulate osmotic pressure, or when a disease damages this mechanism, there is a tendency to accumulate toxic waste and water, which can have dire consequences.

Mammalian systems have evolved to regulate not only the overall osmotic pressure across membranes, but also specific concentrations of important electrolytes in the three major fluid compartments: blood plasma, extracellular fluid, and intracellular fluid. Since osmotic pressure is regulated by the movement of water across membranes, the volume of the fluid compartments can also change temporarily. Because blood plasma is one of the fluid components, osmotic pressures have a direct bearing on blood pressure.

Transport of Electrolytes across Cell Membranes

Electrolytes, such as sodium chloride, ionize in water, meaning that they dissociate into their component ions. In water, sodium chloride (NaCl), dissociates into the sodium ion (Na+) and the chloride ion (Cl–). The most important ions, whose concentrations are very closely regulated in body fluids, are the cations sodium (Na+), potassium (K+), calcium (Ca+2), magnesium (Mg+2), and the anions chloride (Cl–), carbonate (CO3-2), bicarbonate (HCO3–), and phosphate(PO3–). Electrolytes are lost from the body during urination and perspiration. For this reason, athletes are encouraged to replace electrolytes and fluids during periods of increased activity and perspiration.

Osmotic pressure is influenced by the concentration of solutes in a solution. It is directly proportional to the number of solute atoms or molecules and not dependent on the size of the solute molecules. Because electrolytes dissociate into their component ions, they, in essence, add more solute particles into the solution and have a greater effect on osmotic pressure, per mass than compounds that do not dissociate in water, such as glucose.

Water can pass through membranes by passive diffusion. If electrolyte ions could passively diffuse across membranes, it would be impossible to maintain specific concentrations of ions in each fluid compartment therefore they require special mechanisms to cross the semi-permeable membranes in the body. This movement can be accomplished by facilitated diffusion and active transport. Facilitated diffusion requires protein-based channels for moving the solute. Active transport requires energy in the form of ATP conversion, carrier proteins, or pumps in order to move ions against the concentration gradient.

Osmosis

Osmosis is the diffusion of water across a membrane in response to osmotic pressure caused by an imbalance of molecules on either side of the membrane. Osmoregulation is the process of maintenance of salt and water balance (osmotic balance) across membranes within the body’s fluids, which are composed of water, plus electrolytes and non-electrolytes. An electrolyte is a solute that dissociates into ions when dissolved in water. A non-electrolyte, in contrast, doesn’t dissociate into ions during water dissolution. Both electrolytes and non-electrolytes contribute to the osmotic balance. The body’s fluids include blood plasma, the cytosol within cells, and interstitial fluid, the fluid that exists in the spaces between cells and tissues of the body. The membranes of the body (such as the pleural, serous, and cell membranes) are semi-permeable membranes. Semi-permeable membranes are permeable (or permissive) to certain types of solutes and water. Solutions on two sides of a semi-permeable membrane tend to equalize in solute concentration by movement of solutes and/or water across the membrane. As seen in Figure 28.1, a cell placed in water tends to swell due to gain of water from the hypotonic or “low salt” environment. A cell placed in a solution with higher salt concentration, on the other hand, tends to make the membrane shrivel up due to loss of water into the hypertonic or “high salt” environment. Isotonic cells have an equal concentration of solutes inside and outside the cell; this equalizes the osmotic pressure on either side of the cell membrane which is a semi-permeable membrane.

The left part of this illustration shows shriveled red blood cells bathed in a hypertonic solution. Below this, an diagram shows that upper case H subscript 2 baseline upper case O is leaving the red blood cell. The middle part shows healthy red blood cells bathed in an isotonic solution. A diagram below this shows upper H subscript 2 baseline upper O both entering and exiting the cell. And the right part shows bloated red blood cells bathed in a hypotonic solution. One of the bloated cells in the hypotonic solution bursts. A diagram below this shows upper H subscript 2 baseline upper O enterning the cell.
Figure 28.1 Cells placed in a hypertonic environment tend to shrink due to loss of water. In a hypotonic environment, cells tend to swell due to intake of water. The blood maintains an isotonic environment so that cells neither shrink nor swell. (credit: Mariana Ruiz Villareal)

The body does not exist in isolation. There is a constant input of water and electrolytes into the system. While osmoregulation is achieved across membranes within the body, excess electrolytes and wastes are transported to the kidneys and excreted, helping to maintain osmotic balance.

Reading Question #1

The process of maintaining a balance of alt and water (osmotic balance) across membranes within the body’s fluids is known as

a. osmosis

b. thermoregulation

c. osmoregulation

d. hypertonic

 

Osmoregulators and Osmoconformers

Persons lost at sea without any freshwater to drink are at risk of severe dehydration because the human body cannot adapt to drinking seawater, which is hypertonic in comparison to body fluids. Organisms such as goldfish that can tolerate only a relatively narrow range of salinity are referred to as stenohaline. About 90 percent of all bony fish are restricted to either freshwater or seawater. They are incapable of osmotic regulation in the opposite environment. It is possible, however, for a few fishes like salmon to spend part of their life in freshwater and part in seawater. Organisms like the salmon and molly that can tolerate a relatively wide range of salinity are referred to as euryhaline organisms. This is possible because some fish have evolved osmoregulatory mechanisms to survive in all kinds of aquatic environments. When they live in freshwater, their bodies tend to take up water because the environment is relatively hypotonic, as illustrated in Figure 28.2a. In such hypotonic environments, these fish do not drink much water. Instead, they pass a lot of very dilute urine, and they achieve electrolyte balance by active transport of salts through the gills. When they move to a hypertonic marine environment, these fish start drinking seawater; they excrete the excess salts through their gills and their urine, as illustrated in Figure 28.2b. Most marine invertebrates, on the other hand, may be isotonic with seawater (osmoconformers). Their body fluid concentrations conform to changes in seawater concentration. Cartilaginous fishes’ salt composition of the blood is similar to bony fishes; however, the blood of sharks contains the organic compounds urea and trimethylamine oxide (TMAO). This does not mean that their electrolyte composition is similar to that of seawater. They achieve isotonicity with the sea by storing large concentrations of urea. These animals that secrete urea are called ureotelic animals. TMAO stabilizes proteins in the presence of high urea levels, preventing the disruption of peptide bonds that would occur in other animals exposed to similar levels of urea. Sharks are cartilaginous fish with a rectal gland to secrete salt and assist in osmoregulation.

Illustration A shows a fish in a freshwater environment, where water is absorbed through the skin. To compensate, the fish drinks little water and excretes dilute urine. Sodium, potassium and chlorine ions are lost through the skin, and the fish actively transports these same ions into its gills to compensate for this loss. Illustration B shows a fish in a saltwater environment, where water is lost through the skin. To compensate, the fish drinks ample water and excretes concentrated urine. It absorbs sodium, potassium, and chlorine ions through its skin, and excretes them through its gills.
Figure 28.2 Fish are osmoregulators, but must use different mechanisms to survive in (a) freshwater or (b) saltwater environments. (credit: modification of work by Duane Raver, NOAA)

 

CAREER CONNECTION

Dialysis Technician

Dialysis is a medical process of removing wastes and excess water from the blood by diffusion and ultrafiltration. When kidney function fails, dialysis must be done to artificially rid the body of wastes. This is a vital process to keep patients alive. In some cases, the patients undergo artificial dialysis until they are eligible for a kidney transplant. In others who are not candidates for kidney transplants, dialysis is a life-long necessity.

Dialysis technicians typically work in hospitals and clinics. While some roles in this field include equipment development and maintenance, most dialysis technicians work in direct patient care. Their on-the-job duties, which typically occur under the direct supervision of a registered nurse, focus on providing dialysis treatments. This can include reviewing patient history and current condition, assessing and responding to patient needs before and during treatment, and monitoring the dialysis process. Treatment may include taking and reporting a patient’s vital signs and preparing solutions and equipment to ensure accurate and sterile procedures.

Although the kidneys are the major osmoregulatory organ, the skin and lungs also play a role in the process. Water and electrolytes are lost through sweat glands in the skin, which helps moisturize and cool the skin surface, while the lungs expel a small amount of water in the form of mucous secretions and via evaporation of water vapor.

Reading Question #2

Which of the following is most likely an osmoconformer?

a. catfish

b. trout

c. crab

d. bass

 

Kidneys: The Main Osmoregulatory Organ

The kidneys, illustrated in Figure 28.3, are a pair of bean-shaped structures that are located just below and posterior to the liver in the peritoneal cavity. The adrenal glands sit on top of each kidney and are also called the suprarenal glands. Kidneys filter blood and purify it. All the blood in the human body is filtered many times a day by the kidneys; these organs use up almost 25 percent of the oxygen absorbed through the lungs to perform this function. Oxygen allows the kidney cells to efficiently manufacture chemical energy in the form of ATP through aerobic respiration. The filtrate coming out of the kidneys is called urine.

Illustration shows the placement of the kidneys and bladder in a human man. The two kidneys face one another and are located on the posterior side, about halfway up the back. A renal artery and a renal vein extend from the inside middle of each kidney, toward a major blood vessel that runs up the middle of the body. A ureter runs down from each kidney to the bladder, a sac that sits just above the pelvis. The urethra runs down from the bottom of the bladder and through the penis. The adrenal glands are lumpy masses that sit on top of the kidneys.
Figure 28.3 Kidneys filter the blood, producing urine that is stored in the bladder prior to elimination through the urethra. (credit: modification of work by NCI)
Kidney Structure

Externally, the kidneys are surrounded by three layers, illustrated in Figure 28.4. The outermost layer is a tough connective tissue layer called the renal fascia. The second layer is called the perirenal fat capsule, which helps anchor the kidneys in place. The third and innermost layer is the renal capsule. Internally, the kidney has three regions—an outer cortex, a medulla in the middle, and the renal pelvis in the region called the hilum of the kidney. The hilum is the concave part of the bean-shape where blood vessels and nerves enter and exit the kidney; it is also the point of exit for the ureters. The renal cortex is granular due to the presence of nephrons—the functional unit of the kidney. The medulla consists of multiple pyramidal tissue masses, called the renal pyramids. In between the pyramids are spaces called renal columns through which the blood vessels pass. The tips of the pyramids, called renal papillae, point toward the renal pelvis. There are, on average, eight renal pyramids in each kidney. The renal pyramids along with the adjoining cortical region are called the lobes of the kidney. The renal pelvis leads to the ureter on the outside of the kidney. On the inside of the kidney, the renal pelvis branches out into two or three extensions called the major calyces, which further branch into the minor calyces. The ureters are urine-bearing tubes that exit the kidney and empty into the urinary bladder.

The kidney is shaped like a kidney bean standing on end. Two layers, the outer renal fascia and an inner capsule, cover the outside of the kidney. The inside of the kidney consists of three layers: the outer cortex, the middle medulla and the inner renal pelvis. The renal pelvis is flush with the concave side of the kidney, and empties into the ureter, a tube that runs down outside the concave side of the kidney. Nine renal pyramids are embedded in the medulla, which is the thickest kidney layer. Each renal pyramid is teardrop-shaped, with the narrow end facing the renal pelvis. The renal artery and renal vein enter the concave part of the kidney, just above the ureter. The renal artery and renal vein branch into arterioles and venuoles, respectively, which extend into the kidney and branch into capillaries in the cortex.
Figure 28.4 The internal structure of the kidney is shown. (credit: modification of work by NCI)

Reading Question #3

Which of the following statements about the kidney is false?

  1. The renal pelvis drains into the ureter.
  2. The renal pyramids are in the medulla.
  3. The cortex covers the capsule.
  4. Nephrons are in the renal cortex.

Because the kidney filters blood, its network of blood vessels is an important component of its structure and function. The arteries, veins, and nerves that supply the kidney enter and exit at the renal hilum. Renal blood supply starts with the branching of the aorta into the renal arteries (which are each named based on the region of the kidney they pass through) and ends with the exiting of the renal veins to join the inferior vena cava. The renal arteries split into several segmental arteries upon entering the kidneys. Each segmental artery splits further into several interlobar arteries and enters the renal columns, which supply the renal lobes. The interlobar arteries split at the junction of the renal cortex and medulla to form the arcuate arteries. The arcuate “bow shaped” arteries form arcs along the base of the medullary pyramids. Cortical radiate arteries, as the name suggests, radiate out from the arcuate arteries. The cortical radiate arteries branch into numerous afferent arterioles, and then enter the capillaries supplying the nephrons. Veins trace the path of the arteries and have similar names, except there are no segmental veins.

As mentioned previously, the functional unit of the kidney is the nephron, illustrated in Figure 28.5. Each kidney is made up of over one million nephrons that dot the renal cortex, giving it a granular appearance when sectioned sagittally. There are two types of nephrons—cortical nephrons (85 percent), which are deep in the renal cortex, and juxtamedullary nephrons (15 percent), which lie in the renal cortex close to the renal medulla. A nephron consists of three parts—a renal corpuscle, a renal tubule, and the associated capillary network, which originates from the cortical radiate arteries.

Illustration shows the nephron, a tube-like structure that begins in the kidney cortex. Here, arterioles converge in a bulb-like structure called the glomerulus, which is partly surrounded by a Bowmans capsule. Afferent arterioles enter the glomerulus, and efferent arterioles leave. The glomerulus empties into the proximal convoluted tubule. A long loop, called the loop of Henle, extends from the proximal convoluted tubule to the inner medulla of the kidney, and then back out to the cortex. There, the loop of Henle joins a distal convoluted tubule. The distal convoluted tubule joins a collecting duct, which travels from the medulla back into the cortex, toward the center of the kidney. Eventually, the contents of the renal pyramid empty into the renal pelvis, and then the ureter.
Figure 28.5 The nephron is the functional unit of the kidney. The glomerulus and convoluted tubules are located in the kidney cortex, while collecting ducts are located in the pyramids of the medulla. (credit: modification of work by NIDDK)

Reading Question #4

Which of the following statements about the nephron is false?

  1. The collecting duct empties into the distal convoluted tubule.
  2. The Bowman’s capsule surrounds the glomerulus.
  3. The loop of Henle is between the proximal and distal convoluted tubules.
  4. The loop of Henle empties into the distal convoluted tubule.

    Renal Corpuscle

    The renal corpuscle, located in the renal cortex, is made up of a network of capillaries known as the glomerulus and the capsule, a cup-shaped chamber that surrounds it, called the glomerular or Bowman’s capsule.

    Renal Tubule

    The renal tubule is a long and convoluted structure that emerges from the glomerulus and can be divided into three parts based on function. The first part is called the proximal convoluted tubule (PCT) due to its proximity to the glomerulus; it stays in the renal cortex. The second part is called the loop of Henle, or nephritic loop, because it forms a loop (with descending and ascending limbs) that goes through the renal medulla. The third part of the renal tubule is called the distal convoluted tubule (DCT) and this part is also restricted to the renal cortex. The DCT, which is the last part of the nephron, connects and empties its contents into collecting ducts that line the medullary pyramids. The collecting ducts amass contents from multiple nephrons and fuse together as they enter the papillae of the renal medulla.

    Capillary Network within the Nephron

    The capillary network that originates from the renal arteries supplies the nephron with blood that needs to be filtered. The branch that enters the glomerulus is called the afferent arteriole. The branch that exits the glomerulus is called the efferent arteriole. Within the glomerulus, the network of capillaries is called the glomerular capillary bed. Once the efferent arteriole exits the glomerulus, it forms the peritubular capillary network, which surrounds and interacts with parts of the renal tubule. In cortical nephrons, the peritubular capillary network surrounds the PCT and DCT. In juxtamedullary nephrons, the peritubular capillary network forms a network around the loop of Henle and is called the vasa recta.

    LINK TO LEARNING

    Go to this website to see another coronal section of the kidney and to explore an animation of the workings of nephrons.

    Kidney Function and Physiology

    Kidneys filter blood in a three-step process. First, the nephrons filter blood that runs through the capillary network in the glomerulus. Almost all solutes, except for proteins, are filtered out into the glomerulus by a process called glomerular filtration. Second, the filtrate is collected in the renal tubules. Most of the solutes get reabsorbed in the PCT by a process called tubular reabsorption. In the loop of Henle, the filtrate continues to exchange solutes and water with the renal medulla and the peritubular capillary network. Water is also reabsorbed during this step. Then, additional solutes and wastes are secreted into the kidney tubules during tubular secretion, which is, in essence, the opposite process to tubular reabsorption. The collecting ducts collect filtrate coming from the nephrons and fuse in the medullary papillae. From here, the papillae deliver the filtrate, now called urine, into the minor calyces that eventually connect to the ureters through the renal pelvis. This entire process is illustrated in Figure 28.6.

    Illustration labels parts of a nephron and their function. The nephron begins at the glomerulus, a spherical structure that filters small solutes from the blood. The filtrate then enters a winding proximal convoluted tubule, which reabsorbs ions, water, and nutrients, and removes toxins and adjusts the filtrate p H. The proximal convoluted tubule empties into the descending loop of Henle. Aquaporins in the descending loop allow water to pass from the filtrate to the interstitial fluid. The descending loop of Henle turns into the ascending loop of Henle. Both the descending loop and ascending loop are thin at the bottom, and turn thick about a third of the way up. In the ascending loop of Henle, sodium and chlorine ions are reabsorbed from the filtrate into the interstitial fluid. The ascending loop of Henle empties into the distal convoluted tubule, which selectively secretes and absorbs ions to maintain blood pH and electrolyte balance. The distal convoluted tubule empties into a collecting duct, which reabsorbs water and solutes from the filtrate. The collecting duct travels down, toward the middle of the kidney.
    Figure 28.6 Each part of the nephron performs a different function in filtering waste and maintaining homeostatic balance. (1) The glomerulus forces small solutes out of the blood by pressure. (2) The proximal convoluted tubule reabsorbs ions, water, and nutrients from the filtrate into the interstitial fluid, and actively transports toxins and drugs from the interstitial fluid into the filtrate. The proximal convoluted tubule also adjusts blood pH by selectively secreting ammonia (NH3) into the filtrate, where it reacts with H+ to form NH4+. The more acidic the filtrate, the more ammonia is secreted. (3) The descending loop of Henle is lined with cells containing aquaporins that allow water to pass from the filtrate into the interstitial fluid. (4) In the thin part of the ascending loop of Henle, Na+ and Cl- ions diffuse into the interstitial fluid. In the thick part, these same ions are actively transported into the interstitial fluid. Because salt but not water is lost, the filtrate becomes more dilute as it travels up the limb. (5) In the distal convoluted tubule, K+ and H+ ions are selectively secreted into the filtrate, while Na+, Cl-, and HCO3- ions are reabsorbed to maintain pH and electrolyte balance in the blood. (6) The collecting duct reabsorbs solutes and water from the filtrate, forming dilute urine. (credit: modification of work by NIDDK)

    Glomerular Filtration

    Glomerular filtration filters out most of the solutes due to high blood pressure and specialized membranes in the afferent arteriole. The blood pressure in the glomerulus is maintained independent of factors that affect systemic blood pressure. The “leaky” connections between the endothelial cells of the glomerular capillary network allow solutes to pass through easily. All solutes in the glomerular capillaries, except for macromolecules like proteins, pass through by passive diffusion. There is no energy requirement at this stage of the filtration process. Glomerular filtration rate (GFR) is the volume of glomerular filtrate formed per minute by the kidneys. GFR is regulated by multiple mechanisms and is an important indicator of kidney function.

    LINK TO LEARNING

    To learn more about the vascular system of kidneys, click through this review and the steps of blood flow.

    Tubular Reabsorption and Secretion

    Tubular reabsorption occurs in the PCT part of the renal tubule. Almost all nutrients are reabsorbed, and this occurs either by passive or active transport. Reabsorption of water and some key electrolytes are regulated and can be influenced by hormones. Sodium (Na+) is the most abundant ion and most of it is reabsorbed by active transport and then transported to the peritubular capillaries. Because Na+ is actively transported out of the tubule, water follows it to even out the osmotic pressure. Water is also independently reabsorbed into the peritubular capillaries due to the presence of aquaporins, or water channels, in the PCT. This occurs due to the low blood pressure and high osmotic pressure in the peritubular capillaries. However, every solute has a transport maximum and the excess is not reabsorbed.

    In the loop of Henle, the permeability of the membrane changes. The descending limb is permeable to water, not solutes; the opposite is true for the ascending limb. Additionally, the loop of Henle invades the renal medulla, which is naturally high in salt concentration and tends to absorb water from the renal tubule and concentrate the filtrate. The osmotic gradient increases as it moves deeper into the medulla. Because two sides of the loop of Henle perform opposing functions, as illustrated in Figure 28.7, it acts as a countercurrent multiplier. The vasa recta around it acts as the countercurrent exchanger.

    A U-shaped tube represents the loop of Henle. Filtrate enters the descending limb, and exits the ascending limb. The descending limb is water-permeable, and water travels from the limb to the interstitial space. As a consequence, the osmolality of the filtrate inside the limb increases from 300 milliosmoles per liter at the top to 1200 milliosmoles per liter at the bottom. The ascending limb is permeable to sodium and chloride ions. Because the osmolality inside bottom part of the limb is higher than the interstitial fluid, these ions diffuse out of the ascending limb. Higher up, sodium is actively transported out of the limb, and chloride follows. At the top of the ascending limb, the filtrate is 100 milliosmoles.
    Figure 28.7 The loop of Henle acts as a countercurrent multiplier that uses energy to create concentration gradients. The descending limb is water permeable. Water flows from the filtrate to the interstitial fluid, so osmolality inside the limb increases as it descends into the renal medulla. At the bottom, the osmolality is higher inside the loop than in the interstitial fluid. Thus, as filtrate enters the ascending limb, Na+ and Cl- ions exit through ion channels present in the cell membrane. Further up, Na+ is actively transported out of the filtrate and Cl- follows. Osmolarity is given in units of milliosmoles per liter (mOsm/L).

    Loop diuretics are drugs sometimes used to treat hypertension. These drugs inhibit the reabsorption of Na+ and Cl– ions by the ascending limb of the loop of Henle. A side effect is that they increase urination. Why do you think this is the case?

    By the time the filtrate reaches the DCT, most of the urine and solutes have been reabsorbed. If the body requires additional water, all of it can be reabsorbed at this point. Further reabsorption is controlled by hormones, which will be discussed in a later section. Excretion of wastes occurs due to lack of reabsorption combined with tubular secretion. Undesirable products like metabolic wastes, urea, uric acid, and certain drugs, are excreted by tubular secretion. Most of the tubular secretion happens in the DCT, but some occurs in the early part of the collecting duct. Kidneys also maintain an acid-base balance by secreting excess H+ ions.

    Although parts of the renal tubules are named proximal and distal, in a cross-section of the kidney, the tubules are placed close together and in contact with each other and the glomerulus. This allows for exchange of chemical messengers between the different cell types. For example, the DCT ascending limb of the loop of Henle has masses of cells called macula densa, which are in contact with cells of the afferent arterioles called juxtaglomerular cells. Together, the macula densa and juxtaglomerular cells form the juxtaglomerular complex (JGC). The JGC is an endocrine structure that secretes the enzyme renin and the hormone erythropoietin. When hormones trigger the macula densa cells in the DCT due to variations in blood volume, blood pressure, or electrolyte balance, these cells can immediately communicate the problem to the capillaries in the afferent and efferent arterioles, which can constrict or relax to change the glomerular filtration rate of the kidneys.

    CAREER CONNECTION

    Nephrologist

    A nephrologist studies and deals with diseases of the kidneys—both those that cause kidney failure (such as diabetes) and the conditions that are produced by kidney disease (such as hypertension). Blood pressure, blood volume, and changes in electrolyte balance come under the purview of a nephrologist.

    Nephrologists usually work with other physicians who refer patients to them or consult with them about specific diagnoses and treatment plans. Patients are usually referred to a nephrologist for symptoms such as blood or protein in the urine, very high blood pressure, kidney stones, or renal failure.

    Nephrology is a subspecialty of internal medicine. To become a nephrologist, medical school is followed by additional training to become certified in internal medicine. An additional two or more years is spent specifically studying kidney disorders and their accompanying effects on the body.

    Microorganisms and invertebrate animals use more primitive and simple mechanisms to get rid of their metabolic wastes than the mammalian system of kidney and urinary function. Three excretory systems evolved in organisms before complex kidneys: vacuoles, flame cells, and Malpighian tubules.

    Contractile Vacuoles in Microorganisms

    The most fundamental feature of life is the presence of a cell. In other words, a cell is the simplest functional unit of a life. Bacteria are unicellular, prokaryotic organisms that have some of the least complex life processes in place; however, prokaryotes such as bacteria do not contain membrane-bound vacuoles. The cells of microorganisms like bacteria, protozoa, and fungi are bound by cell membranes and use them to interact with the environment. Some cells, including some leucocytes in humans, are able to engulf food by endocytosis—the formation of vesicles by involution of the cell membrane within the cells. The same vesicles are able to interact and exchange metabolites with the intracellular environment. In some unicellular eukaryotic organisms such as the amoeba, shown in Figure 28.8, cellular wastes and excess water are excreted by exocytosis, when the contractile vacuoles merge with the cell membrane and expel wastes into the environment. Contractile vacuoles (CV) should not be confused with vacuoles, which store food or water.

    In this illustration, a cell extends a pseudopod to consume a food particle. The consumed particle is encapsulated in a vesicle. The vesicle fuses with a lysosome, and proteins inside the lysosome digest the food particle. After the food is digested, the vesicle fuses with the cell membrane, and undigested remains are excreted.
    Figure 28.8 Some unicellular organisms, such as the amoeba, ingest food by endocytosis. The food vesicle fuses with a lysosome, which digests the food. Waste is excreted by exocytosis.

    Flame Cells of Planaria and Nephridia of Worms

    As multicellular systems evolved to have organ systems that divided the metabolic needs of the body, individual organs evolved to perform the excretory function. Planaria are flatworms that live in freshwater. Their excretory system consists of two tubules connected to a highly branched duct system. The cells in the tubules are called flame cells (or protonephridia) because they have a cluster of cilia that looks like a flickering flame when viewed under the microscope, as illustrated in Figure 28.9a. The cilia propel waste matter down the tubules and out of the body through excretory pores that open on the body surface; cilia also draw water from the interstitial fluid, allowing for filtration. Any valuable metabolites are recovered by reabsorption. Flame cells are found in flatworms, including parasitic tapeworms and free-living planaria. They also maintain the organism’s osmotic balance.

    Illustration A shows a flame cell, which is bulb-shaped with cilia projecting from one end. The cilia form a point, like the tip of a paintbrush, inside as wide opening at the end of a tube cell. The tube cell narrows into a tubule, then widens into a body where the nucleus is located. The tubule continues past the cell body. Illustration B shows a cross section of an earthworm, which is segmented with walls separating each segment. The trumpet-like opening of a nephridium sticks out of the wall. Cilia surround the opening. Beyond the wall, the nephridium forms a tube that winds down to the ventral surface, where it connects with an opening to the exterior. Just above the opening the tube widens into a bladder.
    Figure 28.9 In the excretory system of the (a) planaria, cilia of flame cells propel waste through a tubule formed by a tube cell. Tubules are connected into branched structures that lead to pores located all along the sides of the body. The filtrate is secreted through these pores. In (b) annelids such as earthworms, nephridia filter fluid from the coelom, or body cavity. Beating cilia at the opening of the nephridium draw water from the coelom into a tubule. As the filtrate passes down the tubules, nutrients and other solutes are reabsorbed by capillaries. Filtered fluid containing nitrogenous and other wastes is stored in a bladder and then secreted through a pore in the side of the body.

    Earthworms (annelids) have slightly more evolved excretory structures called nephridia, illustrated in Figure 28.9b. A pair of nephridia is present on each segment of the earthworm. They are similar to flame cells in that they have a tubule with cilia. Excretion occurs through a pore called the nephridiopore. They are more evolved than the flame cells in that they have a system for tubular reabsorption by a capillary network before excretion.

    Malpighian Tubules of Insects

    Malpighian tubules are found lining the gut of some species of arthropods, such as the bee illustrated in Figure 28.10. They are usually found in pairs and the number of tubules varies with the species of insect. Malpighian tubules are convoluted, which increases their surface area, and they are lined with microvilli for reabsorption and maintenance of osmotic balance. Malpighian tubules work cooperatively with specialized glands in the wall of the rectum. Body fluids are not filtered as in the case of nephridia; urine is produced by tubular secretion mechanisms by the cells lining the Malpighian tubules that are bathed in hemolymph (a mixture of blood and interstitial fluid that is found in insects and other arthropods as well as most mollusks). Metabolic wastes like uric acid freely diffuse into the tubules. There are exchange pumps lining the tubules, which actively transport H+ ions into the cell and K+ or Na+ ions out; water passively follows to form urine. The secretion of ions alters the osmotic pressure which draws water, electrolytes, and nitrogenous waste (uric acid) into the tubules. Water and electrolytes are reabsorbed when these organisms are faced with low-water environments, and uric acid is excreted as a thick paste or powder. Not dissolving wastes in water helps these organisms to conserve water; this is especially important for life in dry environments.

    Illustration shows the digestive tract of a bee. Food enters the mouth, and then goes through the stomach to the intestine. The Malpighian tubules are wormlike protrusions that form a band around the intestine. After the intestine, food enters a bulge called the rectum, and exits through the anus.
    Figure 28.10 Malpighian tubules of insects and other terrestrial arthropods remove nitrogenous wastes and other solutes from the hemolymph. Na+ and/or K+ ions are actively transported into the lumen of the tubules. Water then enters the tubules via osmosis, forming urine. The urine passes through the intestine, and into the rectum. There, nutrients diffuse back into the hemolymph. Na+ and/or K+ ions are pumped into the hemolymph, and water follows. The concentrated waste is then excreted.

    Nitrogenous Waste in Terrestrial Animals: The Urea Cycle

    Of the four major macromolecules in biological systems, both proteins and nucleic acids contain nitrogen. During the catabolism, or breakdown, of nitrogen-containing macromolecules, carbon, hydrogen, and oxygen are extracted and stored in the form of carbohydrates and fats. Excess nitrogen is excreted from the body. Nitrogenous wastes tend to form toxic ammonia, which raises the pH of body fluids. The formation of ammonia itself requires energy in the form of ATP and large quantities of water to dilute it out of a biological system. Animals that live in aquatic environments tend to release ammonia into the water. Animals that excrete ammonia are said to be ammonotelic. Terrestrial organisms have evolved other mechanisms to excrete nitrogenous wastes. The animals must detoxify ammonia by converting it into a relatively nontoxic form such as urea or uric acid. Mammals, including humans, produce urea, whereas reptiles and many terrestrial invertebrates produce uric acid. Animals that secrete urea as the primary nitrogenous waste material are called ureotelic animals.

    The urea cycle is the primary mechanism by which mammals convert ammonia to urea. Urea is made in the liver and excreted in urine. The overall chemical reaction by which ammonia is converted to urea is 2 NH3 (ammonia) + CO2 + 3 ATP + H2O → H2N-CO-NH2 (urea) + 2 ADP + 4 Pi + AMP.

    The urea cycle utilizes five intermediate steps, catalyzed by five different enzymes, to convert ammonia to urea, as shown in Figure 28.11. The amino acid L-ornithine gets converted into different intermediates before being regenerated at the end of the urea cycle. Hence, the urea cycle is also referred to as the ornithine cycle. The enzyme ornithine transcarbamylase catalyzes a key step in the urea cycle and its deficiency can lead to accumulation of toxic levels of ammonia in the body. The first two reactions occur in the mitochondria and the last three reactions occur in the cytosol. Urea concentration in the blood, called blood urea nitrogen or BUN, is used as an indicator of kidney function.

    The urea cycle begins in the mitochondrion, where bicarbonate, shown as upper case H upper case C upper case O subscript 3 baseline is combined with ammonia, shown as upper case N upper case H subscript 3 baseline to make carbamoyl phosphate. Two A T P are used in the process. Ornithine transcarbamylase adds the carbamoyl phosphate to a five-carbon amino acid called ornithine to make L citrulline. L citrulline leaves the mitochondrion, and an enzyme called arginosuccinate synthetase adds a four carbon amino acid called L aspartate to it to make arginosuccinate. In the process, one A T P is converted to A M P and upper case P upper case P lower case i. Arginosuccinate lyase removes a four carbon fumarate molecule from the arginosuccinate, forming the six carbon amino acid L arginine. Arginase 1 removes a urea molecule from the L arginine, forming ornithine in the process. Urea has a single carbon double bonded to an oxygen and single bonded to two ammonia groups. Ornithine enters the mitochondrion, completing the cycle.
    Figure 28.11 The urea cycle converts ammonia to urea.

    EVOLUTION CONNECTION

    Excretion of Nitrogenous Waste

    The theory of evolution proposes that life started in an aquatic environment. It is not surprising to see that biochemical pathways like the urea cycle evolved to adapt to a changing environment when terrestrial life forms evolved. Arid conditions probably led to the evolution of the uric acid pathway as a means of conserving water.

     

    Nitrogenous Waste in Birds and Reptiles: Uric Acid

    Birds, reptiles, and most terrestrial arthropods convert toxic ammonia to uric acid or the closely related compound guanine (guano) instead of urea. Mammals also form some uric acid during breakdown of nucleic acids. Uric acid is a compound similar to purines found in nucleic acids. It is water insoluble and tends to form a white paste or powder; it is excreted by birds, insects, and reptiles. Conversion of ammonia to uric acid requires more energy and is much more complex than conversion of ammonia to urea Figure 28.12.

    Part A shows a photo of a freshwater fish and states that many invertebrates and aquatic species excrete ammonia. The chemical structure of ammonia is upper case N upper case H subscript 3 baseline. Part B shows a photo of a wood rat and states that mammals, many adult amphibians, and some marine species excrete urea. The chemical structure of urea is shown. Urea has one upper N upper H subscript 2 baseline group, and one upper N upper H subscript 2 baseline group attached to a central carbon. An oxygen is also double-bonded to this central carbon. Part C shows a photo of a pigeon and states that insects, land snails, birds, and many reptiles excrete uric acid. The chemical structure of uric acid is shown. Uric acid has a six-membered carbon ring attached to a five-membered ring. Each ring has two upper N upper H groups embedded in it. An oxygen is double-bonded to each ring.
    Figure 28.12 Nitrogenous waste is excreted in different forms by different species. These include (a) ammonia, (b) urea, and (c) uric acid. (credit a: modification of work by Eric Engbretson, USFWS; credit b: modification of work by B. “Moose” Peterson, USFWS; credit c: modification of work by Dave Menke, USFWS)

    EVERYDAY CONNECTION

    Gout

    Mammals use uric acid crystals as an antioxidant in their cells. However, too much uric acid tends to form kidney stones and may also cause a painful condition called gout, where uric acid crystals accumulate in the joints, as illustrated in Figure 28.13. Food choices that reduce the amount of nitrogenous bases in the diet help reduce the risk of gout. For example, tea, coffee, and chocolate have purine-like compounds, called xanthines, and should be avoided by people with gout and kidney stones.

    Photo shows a person's feet. One foot is swollen and red, while the other appears normal.
    Figure 28.13 Gout causes the inflammation visible in this person’s left big toe joint. (credit: “Gonzosft”/Wikimedia Commons)

    Reading Question #5

    The primary mechanism by which mammals convert ammonia to urea is called

    a. nitrogen fixation

    b. nitrogen cycle

    c. urea cycle

     

    Hormones and Kidneys

    While the kidneys operate to maintain osmotic balance and blood pressure in the body, they also act in concert with hormones. Hormones are small molecules that act as messengers within the body. Hormones are typically secreted from one cell and travel in the bloodstream to affect a target cell in another portion of the body. Different regions of the nephron bear specialized cells that have receptors to respond to chemical messengers and hormones. Table 28.1 summarizes the hormones that control the osmoregulatory functions.
    Hormones That Affect Osmoregulation
    Hormone Where produced Function
    Epinephrine and Norepinephrine Adrenal medulla Can decrease kidney function temporarily by vasoconstriction
    Renin Kidney nephrons Increases blood pressure by acting on angiotensinogen
    Angiotensin Liver Angiotensin II affects multiple processes and increases blood pressure
    Aldosterone Adrenal cortex Prevents loss of sodium and water
    Anti-diuretic hormone (vasopressin) Hypothalamus (stored in the posterior pituitary) Prevents water loss
    Atrial natriuretic peptide Heart atrium Decreases blood pressure by acting as a vasodilator and increasing glomerular filtration rate; decreases sodium reabsorption in kidneys
    Table 28.1

    Epinephrine and Norepinephrine

    Epinephrine and norepinephrine are released by the adrenal medulla and nervous system respectively. They are the flight/fight hormones that are released when the body is under extreme stress. During stress, much of the body’s energy is used to combat imminent danger. Kidney function is halted temporarily by epinephrine and norepinephrine. These hormones function by acting directly on the smooth muscles of blood vessels to constrict them. Once the afferent arterioles are constricted, blood flow into the nephrons stops. These hormones go one step further and trigger the renin-angiotensin-aldosterone system.

    Renin-Angiotensin-Aldosterone

    The renin-angiotensin-aldosterone system, illustrated in Figure 28.14 proceeds through several steps to produce angiotensin II, which acts to stabilize blood pressure and volume. Renin (secreted by a part of the juxtaglomerular complex) is produced by the granular cells of the afferent and efferent arterioles. Thus, the kidneys control blood pressure and volume directly. Renin acts on angiotensinogen, which is made in the liver and converts it to angiotensin I. Angiotensin converting enzyme (ACE) converts angiotensin I to angiotensin II. Angiotensin II raises blood pressure by constricting blood vessels. It also triggers the release of the mineralocorticoid aldosterone from the adrenal cortex, which in turn stimulates the renal tubules to reabsorb more sodium. Angiotensin II also triggers the release of anti-diuretic hormone (ADH) from the hypothalamus, leading to water retention in the kidneys. It acts directly on the nephrons and decreases glomerular filtration rate. Medically, blood pressure can be controlled by drugs that inhibit ACE (called ACE inhibitors).

    The renin-angiotensin-aldosterone pathway involves four hormones: renin, which is made in the kidney, angiotensin, which is made in the liver, aldosterone, which is made in the adrenal glands, and A D H, which is made in the hypothalamus and secreted by the posterior pituitary. The adrenal glands are located on top of the kidneys, and the hypothalamus and pituitary are in the brain. The pathway begins when renin converts angiotensinogen into angiotensin I. An enzyme called A C E then converts angiotensin I into angiotensin I I. Angiotensin I I has several direct effects. These include arterial constriction, which increases blood pressure, decreasing the glomerular filtration rate, which results in water retention, and increasing thirst. Angiotensin I I also triggers the release of two other hormones, aldosterone and A D H. Aldosterone causes nephron distal tubules to reabsorb more sodium and water, which increases blood volume. A D H moderates the insertion of aquaporins into the nephridial collecting ducts. As a result, more water is reabsorbed by the blood. A D H also causes arteries to constrict. The hormone A N P is antagonistic to the angiotensin pathway. A N P decreases blood pressure and volume by increasing the glomerulus filtration rate, increasing reabsorption of sodium ions by the nephron, and by inhibiting the release of renin from the kidney and aldosterone from the adrenal gland.
    Figure 28.14 The renin-angiotensin-aldosterone system increases blood pressure and volume. The hormone ANP has antagonistic effects. (credit: modification of work by Mikael Häggström)

    Mineralocorticoids

    Mineralocorticoids are hormones synthesized by the adrenal cortex that affect osmotic balance. Aldosterone is a mineralocorticoid that regulates sodium levels in the blood. Almost all of the sodium in the blood is reclaimed by the renal tubules under the influence of aldosterone. Because sodium is always reabsorbed by active transport and water follows sodium to maintain osmotic balance, aldosterone manages not only sodium levels but also the water levels in body fluids. In contrast, the aldosterone also stimulates potassium secretion concurrently with sodium reabsorption. In contrast, absence of aldosterone means that no sodium gets reabsorbed in the renal tubules and all of it gets excreted in the urine. In addition, the daily dietary potassium load is not secreted and the retention of K+ can cause a dangerous increase in plasma K+ concentration. Patients who have Addison’s disease have a failing adrenal cortex and cannot produce aldosterone. They lose sodium in their urine constantly, and if the supply is not replenished, the consequences can be fatal.

    Antidiuretic Hormone

    As previously discussed, antidiuretic hormone or ADH (also called vasopressin), as the name suggests, helps the body conserve water when body fluid volume, especially that of blood, is low. It is formed by the hypothalamus and is stored and released from the posterior pituitary. It acts by inserting aquaporins in the collecting ducts and promotes reabsorption of water. ADH also acts as a vasoconstrictor and increases blood pressure during hemorrhaging.

    Atrial Natriuretic Peptide Hormone

    The atrial natriuretic peptide (ANP) lowers blood pressure by acting as a vasodilator. It is released by cells in the atrium of the heart in response to high blood pressure and in patients with sleep apnea. ANP affects salt release, and because water passively follows salt to maintain osmotic balance, it also has a diuretic effect. ANP also prevents sodium reabsorption by the renal tubules, decreasing water reabsorption (thus acting as a diuretic) and lowering blood pressure. Its actions suppress the actions of aldosterone, ADH, and renin.

     

    Adapted from Clark, M.A., Douglas, M., and Choi, J. (2018). Biology 2e. OpenStax. Retrieved from https://openstax.org/books/biology-2e/pages/1-introduction

     

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