9. Homeostasis 9.1 Concept of homeostasis Definition Homeostasis is the maintenance of a constant internal environment within a living organism.

(homoio means the same; stasis means standing) The internal environment of a multicellular organism is the tissue fluid bathing the cells. Conditions that are kept constant include: 1) pH 2) blood pressure 3) temperature 4) salt content 5) blood glucose Importance The enzymes that control the chemical reactions in our cells operate best within fairly narrow limits of temperature, pH, substrate and product concentration. So for cells to survive and function efficiently, it is important that the composition of tissue fluid be kept as stable as possible. Control mechanisms and feedback mechanisms Each homeostatic control system must have: 1. a receptor/detector (sensor), which detects a stimulus. A stimulus is a change in the level of the factor being regulated. This detectable change is called the input. 2. a coordinator/controller, which receives and coordinates information from the receptor and sends out instructions which triggers the action that will correct the deviation 3. an effector, which carries out the action that brings about the necessary change needed to return the system its original level (often called the corrective mechanism) 4. a feedback loop, which informs the receptor of any change in the system as a result of action by the effector Negative feedback Positive feedback

Negative feedback Homeostasis is achieved by a process called negative feedback. A change in the level of an internal factor causes effectors to restore the internal environment to its original level. For example, an increase in the internal body temperature causes the body to lose more heat; a decrease in body temperature causes the body to generate more heat. This type of system, in which a change in the level of a factor triggers a corrective mechanism, is called a self-adjusting system.

Positive feedback Sometimes a homeostatic mechanism breaks down and negative feedback does not occur. Deviations from the set point are not corrected but made even larger. This result in a process called positive feedback in which a small change in output causes further change in the same direction. Positive feedback is usually harmful because it tends to produce unstable conditions. For example, when the negative feedback mechanisms in mammalian temperature regulation break down, a rise in body temperature can spiral upwards and threaten death. However, in certain circumstances positive feedback can be useful as in oxytocin release during childbirth.

Temperature regulation 1. The temperature of environments inhabited by living organisms ranges from 90 C in hot springs to 40 C in the Arctic. 2. Most organisms, however, live in the narrow range of temperature 0-30 C. To survive, most animals need to exert some control over their body temperature. This regulation of body temperature is called thermoregulation. 3. In all organisms heat may be gained in two main ways: a) metabolism of food b) absorption of solar energy 4. Heat may be lost in four main ways: a) evaporation of water, e.g. during sweating c) conduction from the body to the ground or other objects b) convection from the body to the air or water d) radiation from the body to the air, water or ground. 5. Animals that can maintain a stable body temperature independently of their environment are termed endotherms (sometimes called homoiotherms or warm blooded), e.g. all mammals and birds. 6. Animals with body temperature that changes with the environmental temperature (for example, most animals except birds and mammals, are termed ectotherms (sometimes called poikilotherms or cold blooded). Thermoregulation in ectotherms 1. Ectotherms are animals that do not generate much body heat. The body temperature frequently fluctuates with that of the environment, such as fish and amphibians. 2. Ectotherms do attempt to regulate their temperatures within broad limits by their behaviour or by increased activity, such as lizards. 3. Many lizards live in habitats that are hot during the day but cold at night. In the morning and evening, it basks in the sun to raise its body temperature so that it is fully active and able to hunt. During the hottest part of the day, the lizards find shade under a rock to prevent itself from overheating. At night, they live in a burrow and its body temperature again drops to nearer that of its surroundings. Thermoregulation in endotherms 1. Endotherms are animals that generate their own body heat. They are able to regulate their body temperature and keep it relatively constant, usually in the range of 35 - 44 C, using physiological, anatomical as well as behavioural adaptations. 2. The higher the body temperature, the higher the metabolic rate of the animal. This is especially important for birds during flight where the energy demand is high; therefore they have body temperatures in the range of 40 44 C.

Control of body temperature 1. Body temperature is controlled by the hypothalamus, a small structure at the base of the midbrain just above the pituitary gland. It contains the thermoregulatory centre which has two parts: a heat gain centre and a heat loss centre. Core body temperature stays at around 37 C. 2. The hypothalamus monitors the temperature of blood passing through it (the detectors are thermoreceptors) and in addition receives nervous information from hot and cold receptors in the dermis of the skin about external temperature changes. 3. Any reduction in blood temperature will bring about changes which conserve heat. The hypothalamus sends out nerve impulses that switch on warming mechanisms. A rise in blood temperature has the opposite effect. The hypothalamus sends out nerve impulses that switch on cooling mechanisms, such as increased sweating and vasodilation. Control of blood glucose level The pancreas The pancreas is both an exocrine and an endocrine gland. Acinar cells (exocrine cells) in the pancreas secretes pancreatic juice down the pancreatic duct into the duodenum. Throughout the pancreas are patches of endocrine tissues called islets of Langerhans. The islets contain two types of cells: -cells, which produce the hormone glucagon, and -cells, which produce the hormone insulin. The hormones are discharged directly into the blood and they play a central role in the control of blood glucose.

Effects of insulin and glucagon on the levels of blood glucose When the blood glucose concentration falls below the set point, the alpha cells (-cells) secrete the hormone glucagon, a single polypeptide chain with 29 amino acids. Glucagon activates the enzyme phosphorylase in the liver and muscles which catalyses the breakdown of glycogen to glucose (glycogenolysis). Glucagon also increases the conversion of amino acids and glycerol into glucose-6-phosphate (gluconeogenesis). The smaller beta cells (-cells) detect increases in blood glucose levels above the normal and secrete a different hormone called insulin (2 polypeptide chains of 51 amino acids). Insulin causes excess glucose to become converted to glycogen in liver and muscle cells (glycogenesis). Other changes include an increase in the rate of respiration in the cells with more available glucose and an increase in the rate at which glucose is converted to fat and stored in adipose tissue.

1. In addition to glucagons, at least 2 other hormones can also increase blood glucose concentrations. 2. Adrenaline is secreted in times of acute stress or excitement. It causes the breakdown of glycogen in the liver, boosting blood glucose concentrations. 3. When glycogen stores in the liver become exhausted, cortisol is secreted by the adrenal glands. Cortisol promotes liver cells to convert amino acids and glycerol into glucose.

Fall in blood glucose level

Rise in blood glucose level

Diabetes mellitus 1. If the pancreas cannot secrete sufficient insulin, or if target cells lose their responsiveness to insulin, the blood glucose concentration can reach dangerously high levels. The resulting condition is called diabetes mellitus. It can result from incomplete development of, damage to, or disease of the islets of Langerhans in the pancreas which secretes insulin. 2. The blood sugar concentration becomes so high that the kidney is unable to reabsorb all the glucose filtered into its tubules back into the blood. Consequently glucose is excreted in the urine. 3. In diabetics there is insufficient insulin to increase the permeability of the cell membranes to glucose, so the cells are starved of their fuel, even though the blood glucose level is high. The cells have to respire using proteins and fats instead, which results in the patient losing weight.

Glucose tolerance curves blood glucose levels after administration of 50 g of glucose to different individuals.

4. The high blood glucose is called hyperglycaemia, and the presence of glucose in the urine is called glycosuria; both are signs of diabetes mellitus. Other diagnostic features include a lack of energy, a craving for sweet foods, and persistent thirst. 5. The main diagnostic test is a glucose tolerance test in which the patient swallows a sugar solution and the doctor measures the blood glucose concentration at intervals. Graphs of the results from a diabetic and a normal person are distinctly different as seen in the diagram. 6. Some diabetics can control their blood glucose levels by carefully regulating their carbohydrate intake. Others may need daily injections of specific amounts of insulin. 7. For many years, this insulin came from slaughtered animals but now, it is produced on a large scale by genetic engineering, eliminating the problems associated with animal-derived insulin.

Calculation of pressure in movement of fluid between blood

Two forces control movement of fluid through capillary walls. (i) (ii) Hydrostatic pressure which pushes fluids out of capillaries Osmotic pressure which causes water molecules to move back into capillaries.

When HP > OP, there is a net filtration pressure which causes fluids to leave capillaries. When OP > HP, there is a net absorption pressure which causes water to reenter capillaries. STPM (Past year questions)

A 6 mm Hg

B 7 mm Hg

C 13 mm Hg

D 20 mm Hg

E 43 mm Hg

Further examples of questions: STPM 2004 No 17 Topical Practice Exercises (Biology) Vol 1, pg 56. 9.2 Liver Structure and functions in mammals Structure of liver Liver lobules Liver cells in part of a lobule

1. The liver is the largest organ in the human abdomen. It is made up of lobules. In each lobule, blood from the hepatic portal vein and hepatic artery spreads out and flows slowly through sinusoids (blood-filled spaces) past each hepatocyte (liver cell). 2. Hepatocytes have an exceptionally high density of mitochondrias to provide sufficient ATP for all the energyconsuming functions, and microvilli to increase the surface area for uptake of substances from the blood. 3. The liver acts as the bodys chemical-processing factory, carrying out many functions including: a) Storage of minerals and vitamins The liver stores minerals, e.g. iron, potassium, copper and zinc and the vitamins A, D, E, K (fat-soluble) and also B12 and C(water-soluble) which can later be released if deficient in the diet. b) Synthesis of plasma proteins The liver is responsible for the production of vital proteins found in blood plasma such as albumins and globulins as well as the clotting factors prothrombin and fibrinogen) c) Detoxification of poisons - The liver has a group of enzymes which break these chemicals down into less harmful products for example, the enzyme catalase breaks down hydrogen peroxide to water and oxygen. d) carbohydrate metabolism - The liver helps to regulate blood glucose levels; glucose can be added to the blood by converting glycogen to glucose (glycogenolysis), or by converting amino acids and glycerol into glucose (gluconeogenesis); glucose can be removed from the blood by storing the glucose as glycogen (glycogenesis), converting to fat, or using glucose as a fuel for cellular respiration. e) Fat metabolism - Liver cells process fatty acids so that they can be transported and deposited in the body. The liver also regulates the amounts of phospholipids and cholesterol, synthesizing them or eliminating them as required. Excess cholesterol and phospholipids are eliminated in the bile. f) Production of bile Bile is a thick yellow liquid containing water, bile salts, bile pigments, inorganic ions, and cholesterol. Bile salts and bile acids help to break down fat into small droplets (emulsification) which provides a larger surface area for digestion. The liver produces bile salts and adds to them the bile pigment bilirubin from the breakdown of red blood cells. Bile is transported via bile canaliculi (small channels in the liver lobules) to the bile duct which is stored in the gall bladder before being discharged into the duodenum. g) Protein metabolism - Proteins are not stored by the body and so excess amino acids are broken down in the liver by a process called deamination. The amino group is removed and coverted to the less toxic urea in mammals in the ornithine cycle. Transamination reactions, wherby one amino acid is converted to another, are also performed by the liver. All non-essential amino acids may be synthesized in this way, should they be temporarily deficient in the diet. h) Storage and breakdown of red blood cells The liver with its vast complex of blood vessels, forms a large store of blood with a capacity of up to 1500 cm3. The Kupffer cells (star shaped white blood cells) lining the sinusoids break down red blood cells at the end of their 120-day life span, producing the bile pigment bilirubin which is excreted in the bile. The iron is either stored in the liver or used in the formation of new red blood cells by the bone marrow. Cori cycle The process by which the lactic acid produced during exercise is converted to glucose and returned to the muscle is the Cori cycle. During heavy exercise lactic acid is released by muscle cells into the bloodstream. Some lactic acid is taken by the liver and resynthesized to glucose. The resulting glucose may be taken up by muscles that have depleted their glycogen reserves. The oxygen used by the liver to convert lactic acid to glucose is the oxygen debt. Other tissues that respire oxidatively during exercise such as the heart and kidney can convert lactic acid to pyruvate and metabolize it to CO2 by way of the Krebs cycle.

Excreting nitrogenous waste 1. Excess amino acidss are not stored by the body and are broken down in the liver by deamination, a process that involves removal of the amino group (-NH2) to form ammonia (NH3). The remains of the acid molecules can be converted to fats and carbohydrates that can be used immediately or stored. 2. Ammonia is very toxic and very soluble; only aquatic animals can use it as an excretory product as it requires great dilution during their excretion (ammoniotelic). Terrestrial animals convert ammonia to a less toxic and less soluble substances such as urea in mammals (ureotelic) or uric acid in insects, reptiles and birds (uricotelic). Both these forms of excretory products avoid harmful effects to the organisms and help to conserve water. 3. Urea is much less toxic than ammonia and, although it is less soluble, less water is needed for its elimination because the tissues can tolerate higher concentrations of it. Uric acid is non-toxic and almost insoluble in water. It requires very little water for its removal and is therefore a very suitable product for animals living in arid conditions. This is an advantage to flying organisms such as insects and birds as its storage within them does not greatly increase their mass. 4. Mammals convert ammonia to urea by combining two molecules of ammonia with one of carbon dioxide in a series of enzyme-catalysed reactions called the ornithine cycle. Urea is transported in the blood to the kidneys where it is excreted in the urine.

Ornithine cycle Ammonia produced by deamination is converted in the liver into the soluble product urea by a cyclic reaction known as the ornithine cycle. Enzyme arginase in the liver catalyses the synthesis of urea from the amino acid arginine. arginase Arginine + water urea + ornithine Ornithine is converted to citrulline by combining with ammonia and CO2 with the loss of water. Ornithine + ammonia + carbon dioxide citrulline + water Arginine is reformed when citrulline combines with NH3 with the loss of water. Citrulline + ammonia arginine + water Arginine can be hydrolysed to urea and ornithine and the cycle repeats.

9.3.1 Urine formation in animals 1. The kidneys are paired organs concerned with excretion and homeostasis (osmoregulation). They remove nitrogenous waste products, and they also help to control the water content and pH of the blood. 2. Each kidney consists of three main areas; a dark outer region called the cortex, the lighter inner region called the medulla, and a central space called the pelvis. 3. There are about a million nephrons in each human kidney. Nephrons are very small thin-walled tubules between 2 and 4 cm long. At one end of the nephron, in the cortex, is the cup-shaped Bowmans capsule which encloses a small group of capillaries called a glomerulus. Together they are known as a Malpighian body. 4. The capsule leads into a coiled structure called the proximal convoluted tubule. This leads into the long, hairpinlike loop of Henle, which runs deep into the medulla (descending limb) and then back out to the cortex (ascending limb). 5. This leads into a second coiled tube, the distal convoluted tubule which then joins a collecting duct that carries urine through the medulla to the pelvis of the kidney. 6. Blood is brought to the kidney by the renal artery, which branches many times to form arterioles. Each Bowmans capsule is supplied with blood by an afferent arteriole which branches inside the capsule to form a knot of capillaries called the glomerulus. These join up again to form the efferent arteriole, which takes blood away from the capsule and leaves by the renal vein. 7. There are 2 types of nephrons depending on the length and nature of their loops of Henle. Cortical nephrons have shorter loops while juxtamedullary nephrons have longer loops. In humans, about 85% of the nephrons are cortical and 15% are juxtamedullary while nearly all the nephrons have very long loops in some desert-living rodents.

Functions of the nephron The dual function of excretion and osmoregulation of the kidneys can be seen in the functioning of the nephrons. The main stages are: 1. Ultrafiltration in the Bowmans capsule 2. Selective reabsorption of useful substances 3. Tubular secretion of waste substances Ultrafiltration in the Bowmans capsule

1. Ultrafiltration is the separation of large molecules from small ones by a very fine filter. This process takes place in the glomerulus, a knot of capillaries surrounded by the Bowmans capsule. It is a passive process as substances passing from the blood into the glomerular filtrate is made entirely according to size. 2. The capillary wall or endothelium of the glomerulus is a single layer of cells with thousands of gaps (0.1 m in diameter). The endothelium is closely pressed against the basement membrane of the capsule, a thin porous membrane of squamous cells which is the only effective barrier between the blood in the glomerulus and the cavity of the capsule. The inner, or v isceral, layer of the Bowmans capsule has special cells called podocytes with numerous foot-like processes. 3. Blood enters the glomerulus at high pressure through the afferent arteriole from the renal artery. The pressure is kept high because the afferent arteriole has a larger diameter than the efferent arteriole. 4. As a result of this pressure, substances are forced through the endothelial pores of the capillary, across the basement membrane, and down through the spaces between the podocytes into the Bowmans capsule by ultrafiltration. The endothelium and podocytes act as coarse filters, but the basement membrane acts as the fine filter, preventing the passage of large molecules into the nephron. 5. The glomerular filtrate has a solute composition similar to that of blood (e.g. glucose, amino acids, vitamins, some hormones, urea, uric acid, creatinine, ions and water), but it normally lacks red blood cells and plasma proteins which are too large to pass through the basement membrane. 6. Ultrafiltration is a passive process and selection of substances from the blood into the glomerular filtrate is made entirely according to relative molecular mass. 7. The filtrate contains valuable substances and fluids too precious to lose in the urine. Useful substances are selectively reabsorbed back into the body. Without selective reabsorption humans would produce 180 dm3 of urine per day whereas the actual volume produced is approximately 1.5 dm3.

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Selective reabsorption Both passive and active processes are involved in the selective reabsorption of substances from the nephron. The composition is further altered by the active secretion of substances from the blood into the tubule.

A. Proximal convoluted tubule 1. Over 80% of the glomerular filtrate is reabsorbed here. The single layer epithelial cells have special adaptations for reabsorption. The cell surface facing the lumen has numerous microvilli forming a brush border which greatly enlarge the surface area.The base of each cell is adjacent to a blood capillary with numerous intercellular spaces and contains large numbers of mitochondria, providing the ATP necessary for active transport. 2. All the glucose, amino acids, vitamins, hormones and many sodium and chloride ions are reabsorbed from the proximal convoluted tubule by active transport back into the blood. However, if the blood glucose concentration is higher than a critical level called the renal threshold, glucose appears in the urine (diabetes mellitus). 3. About 85% of the water is reabsorbed passively by osmosis. Small protein molecules that have managed to squeeze their way into the tubule are reabsorbed in pinocytotic vesicles. 4. Surprisingly, about 55% of the urea also moves out of the nephron into the tissue fluid and blood. The 45% left in the tubule is excreted in the urine. 5. By the time the fluid leaves the proximal convoluted tubule, the tubular filtrate is isotonic with blood in the surrounding capillaries. Selective reabsorption continues for water and salt along the rest of the nephron, according on the state of the person. B. Loop of Henle 1. The loop of Henle appears to function as a countercurrent exchange mechanism that increases the solute concentration in the medulla of the kidney. By doing so, it creates an osmotic gradient along which water can be withdrawn from the collecting duct by osmosis if circumstances demand it. A concentrated urine can be produced as a result. 2. The ascending limb is more permeable to salts and impermeable to water. As the filtrate moves up, sodium and chloride ions are pumped actively out of the ascending limb into the surrounding tissue fluids. 3. This creates an osmotic gradient which draws water out of the descending limb (highly permeable to water) into the vasa recta (blood vessels) which is both slow-flowing and freely permeable, two factors which aid the uptake of water. As a result the filtrate becomes more concentrated as it passes down the descending limb of the loop.

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4. The net result is that the solute concentration at any part of the loop is lower than it is in the descending limb. The process continues down the length of the descending loop so that this concentrating effect is multiplied. This mechanism is called the countercurrent multiplier mechanism. 5. The countercurrent multiplier means that the fluid in and around the loop of Henle becomes saltier as it goes down the loop, is saltiest at the hairpin bend (tip of the loop) and becomes less salty as it goes up the ascending limb (ion are removed). 6. For animals of the same size, the final salt concentration that can be produced in the tissue fluid depends on the length of the loop: the longer the loop, the higher the final salt concentration. 7. As the collecting ducts pass through the medulla to the pelvis, they pass through this region of high solute concentration, so water is drawn out of the collecting ducts by osmosis, resulting in a far more concentrated urine. C. Distal convoluted tubule 1. The cells in this region are very similar to those of the proximal convoluted tubule, having a brush border and numerous mitochondria. 2. The permeability of their membranes to water and salt is affected by hormones, so reabsorption is dependent on the amount of water and salts present in the body. D. Collecting duct 1. The permeability of the walls of the collecting duct, like the permeability of the distal convoluted tubule, is affected by hormones. This regulates how much water passes out into the medulla. 2. This hormonal effect, together with the hypertonic interstitial fluids built up by the loop of Henle in the medulla, determine whether hypotonic or hypertonic urine is released from the kidney. 3. If the walls of the collecting duct are water-permeable, water leaves the ducts to pass into the hyperosmotic surroundings and into the vasa recta and concentrated urine is produced. 4. If the ducts are impermeable to water the final urine will be less concentrated. 3. Tubular or direct secretion 1. About 80% of the blood plasma which enters the human kidneys is not filtered from the glomeruli into the renal capsules. 2. However, some of the substances like uric acid, ammonium ions, creatinine and certain drugs in the blood may be discharged into the nephrons by direct secretion. This active removal of substances is called tubular secretion. 3. Direct secretion enables greater quantities of such wastes to be eliminated than by ultrafiltration alone. 4. The opposite of reabsorption 5. Molecules are transported out of the peritubular capillaries, through tubule cells, and into the filtrate 6. Eliminates H+ ions, metabolites, and toxins

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Osmoregulation by the kidney - role and mechanism of action of antidiuretic hormone (ADH)

1. Osmoregulation is the homeostatic control of body water. The receptors for detecting changes are osmoreceptors located in the hypothalamus of the brain. They monitor the water potential of the blood as it flows through the hypothalamus. 2. The rise in blood osmotic pressure (low water potential) is detected by osmoreceptors and results in nerve impulses passing to the posterior pituitary gland which releases ADH. ADH increases the permeability of the walls of the distal convoluted tubules and collecting ducts to water, allowing more water to be reabsorbed into the blood by osmosis. ADH also increases the permeability of the collecting duct to urea which passes into the medulla, increasing the osmotic concentration and causing more water to be lost from the descending loop of Henle. Smaller volumes of more concentrated urine are produced and water is conserved in the body. 3. If the osmotic pressure of the blood falls (water potential is higher than normal), less ADH is released from the posterior part of the pituitary gland. The walls of the distal convoluted tubules and collecting ducts remain impermeable to water and urea. Less water is reabsorbed into the medulla and larger quantities of dilute urine are produced. 4. Anyone who is unable to produce sufficient levels of ADH will produce large volumes of very dilute urine, whatever their diet. This condition is termed diabetes insipidus. Control of blood sodium ions by aldosterone 1. Aldosterone is the hormone responsible for maintaining a more or less constant sodium level in the plasma and it has a secondary effect on water reabsorption. 2. If the sodium ion concentration in the blood is low, then the blood water potential increases and water moves by osmosis into the tissues. This results in a slight lowering of blood pressure, monitored by a group of secretory cells (known as juxtaglomerular complex) lying between the afferent arteriole and the distal convoluted tubule to release the enzyme renin. 3. Renin causes a plasma globulin produced by the liver to form the hormone angiotensin. Angiotensin stimulates the release of aldosterone from the adrenal cortex.

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4. Aldosterone causes sodium ions to be actively taken up from the distal convoluted tubule and collecting ducts to the tissue fluid and blood. Water follows by osmosis, and the blood pressure rises until it returns to the set point, thus restoring the sodium level of the plasma and the volume of the blood. 5. If the blood pressure is higher than the set point, negative feedback reduces the secretion of aldosterone.

Control of pH 1. The distal convoluted tubule also helps regulate blood pH, maintaining it at 7.4 by varying the pH of urine. 2. The cells of the tubule combine water and carbon dioxide (produced by the tubule cells as they respire) to form carbonic acid. This then dissociates into hydrogen ions and hydrogen carbonate ions. 3. If the blood pH is too low, more hydrogen ions (protons) are excreted into the lumen of the tubule while the absorption of hydrogen carbonate ions into the blood raises its pH. In the lumen, hydrogen phosphate ions (HPO42-) combine with these hydrogen ions to form dihydrogen phosphate ions (H2PO4-) which are excreted in the urine. The acidic effect of the hydrogen ions is therefore buffered by these hydrogen phosphate ions. 4. If the blood pH is too high, more hydroxide ions are excreted. By these processes, the pH of urine may be as low as 4 or as high as 9. 9.3.2 Plants Role of stomata in the regulation of water loss refer to topic 7.2.1 (Stomata) Adaptation of plants to the environment Some of the most striking adaptations of plants relate to the availability of water in their environment. In fact, plants are often classified into four groups according to their water supply. 1. Mesophytes are plants living in areas where water is readily available 2. Xerophytes live in areas where water is in short supply 3. Hydrophytes are plants in freshwater or very wet environments 4. Halophytes occupy salt marshes, estuarine muds, or areas close to the sea, where the high concentration of salts makes obtaining water difficult

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Mesophytes 1. In hot conditions, mesophytes may begin to overheat and suffer temperature stress. In well watered soils, they can respond to this by increasing their rate of transpiration which cools the plant. 2. In dry conditions, mesophytes often suffer water stress, losing more water to the air than they can regain from the soil. They respond to water stress by closing their stomata. Xerophytes - adaptations to reduce water loss Transverse section through a leaf of marram grass Prickly pear cactus

1. Avery thick waxy cuticle cuts down evaporation from the upper epidermis, as in the leaves of evergreen shrubs such as Oleander. 2. Having smaller leaves reduces the surface area/volume ratio, so there is less area over which water is lost, as in Pinus. 3. Rolling up of leaves so that the lower surface faces inside and traps humid air next to the stomata reduces the rate of evaporation, as in marram grass (Ammophila). 4. Sunken stomata can be found in grooves in some xerophytic leaves, foe example marram. Humid air accumulates in the grooves above the stomata, reducing the rate of diffusion of water molecules. Sunken stomata have the effect of reducing the amount of air movement over the surface of the stomata. 5. Leaf hairs are outgrowths of the epidermal cells of leaves. They are also able to trap damp air close to the leaf surface, reducing the amount of air movement and cutting down transpiration, as in marram and Oleander. 6. Some plants have succulent leaves in which they are able to store water, for example Bryophyllum. 7. Other plants have succulent stems for water storage, for example cacti. These stems also take over the role of photosynthesis from the reduced leaves. 8. Some plants have the ability to close their stomata during daylight to significantly reduce transpiration, for example most cacti and pineapple. 9. Many xerophytes have shallow, extensive root systems to quickly absorb water from rain and overnight condensation, for example most cacti. 10. The development of sclerenchyma tissue in the leaf prevents it from collapsing in times of drought, as in the case of Hakea. 11. An ability to fix carbon dioxide at night so that the stomata can be closed during the day (CAM).

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Hydrophytes

1. The bodies of hydrophytes (water plants) are usually supported by water so they have only small amounts of mechanical and supporting tissues. They have little or no lignified tissue, the xylem is poorly developed, and the root system is either absent or reduced. 2. Hydrophytes have specialized leaves that may be floating, as in water lilies, or finely divided, as in Canadian pondweed (Elodea). 3. Hydrophyte stems and leaves often store gases in large interconnecting air spaces; this enables them to obtain sufficient oxygen and carbon dioxide from water that is not aleays fully aerated. As well as storing gases, these spaces make hydrophytes buoyant when submered. Halophytes 1. Although halophytes (salt plants) such as the glasswort commonly live in waterlogged estuarine muds and slat marshes, they are faced with a water supply problem: the surrounding water is often very salty. They overcome this problem by actively absorbing salts into their roots. As a result, the roots have a lower water potential than the surrounding water, and water can flow into the plant by osmosis in the usual way. 2. Many halophytes have evolved xeromorphic features which help them conserve water. Xeromorphic features of the glasswort include a thick epidermis, waxy cuticle, and a reduced number of stomata.

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