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"Just as we live in a constantly changing world, so do the cells and tissues survive in a constantly changing microenvironment.”

• property of a system that regulates its internal environment & tends to maintain a stable, constant condition of properties like temperature or pH • Internal – constant supply of materials and waste prdts should be expelled • External – response to change
Walter Bradford Cannon coined “homeostasis”

• receptor - component that monitors and responds to changes in the environment - When the receptor senses a stimulus, it sends information to a • control center - component that sets the range at which a variable is maintained - determines an appropriate response to the stimulus - In most homeostatic mechanisms, control center is the brain - sends signals to an • Effector - which can be muscles, organs or other structures that receive signals from the control center. • After receiving the signal, a change occurs to correct the deviation by either enhancing it with positive feedback or depressing it with negative feedback.

Control mechanisms : 3 interdependent components

Positive feedback
• mechanisms designed to accelerate /enhance the output created by a stimulus that has already been activated • push levels out of normal ranges • beneficial but is rarely used by the body due to risks of the acceleration's becoming uncontrollable

Examples: • blood platelet accumulation, which, in turn, causes blood clotting in response to a break or tear in the lining of blood vessels • release of oxytocin to intensify the contractions that take place during childbirth

Negative feedback
• mechanisms consist of reducing the output or activity of any organ or system back to its normal range of functioning Example • when the body is deprived of food • body would reset the metabolic set point to a lower than normal value • Body functions at a slower rate, even though the body is starving. • Therefore, people who deprive themselves of food while trying to lose weight would find it easy to shed weight initially and much harder to lose more after. • due to the body readjusting itself to a lower metabolic set point to allow the body to survive with its low supply of energy. Exercise can change this effect by increasing the metabolic demand.

Water and Osmotic Regulation
How Marine Invertebrates Meet Problems of Salt and Water Balance
• Most marine invertebrates are in osmotic equilibrium with their seawater environment. • body surfaces permeable to water and salts, the internal and external concentrations are equal. • Such animals that cannot regulate osmotic pressure of their body fluids are called osmotic conformers. • This functions for open ocean organisms because the open ocean is stable. • Animals that must live within a narrow salinity range are stenohaline. • Organisms that can tolerate the wide variations found in estuaries are euryhaline. • A hyperosmotic regulator maintains body fluids in higher concentration than surrounding water.

• Kidneys, or antennal glands in a crab, can maintain a higher concentration by excreting excess water. • Salt-secreting cells in the gills that remove ions from seawater counter loss of salt ions. • Any process that requires an expenditure of energy is an active transport process. • Any process that works against the diffusion gradient will require active transport.

• 450M years ago, jawed fishes began to penetrate brackish and freshwater rivers. • unexploited habitat with abundant food presented a physiological evolutionary challenge. • Freshwater fishes must prevent salt loss and unload excess water.

Invasion of Fresh Water : Salt & water balance in freshwater fishes

• The scaled and mucus-covered surface of a fish is nearly waterproof. • Water that enters by osmosis across the gills is pumped out by the kidney as very dilute urine. • Salt-absorbing cells in the gills move sodium and chloride ions from water to blood. • Salt present in the fish’s food also replaces any salt that is lost by diffusion.

• Amphibians that live in water use their skin to transport sodium and chloride; these tissues are standard laboratory models for ion transport

• Marine bony fishes maintain salt concentration of body fluids at about one-third that of seawater. • They are therefore hypoosmotic regulators, maintaining their body fluids at lower concentration. • Modern oceanic bony fishes are descendants of freshwater fishes • Their ionic body concentration of about one-third that of seawater is related to their marine heritage. • Freshwater fish returning to the sea in the Triassic period lost water and gained salt-it “dried out.”

Return of Fishes to the Sea: Salt & water balance in marine fishes

• Marine fish therefore drink seawater and it is absorbed in the intestine; salt is carried by blood to gills. • Special salt-secreting cells in the gills transport the salt back to the sea. • Ions that remain in the intestine as residue (e.g. magnesium, sulfate, calcium) are voided with feces. • Elasmobranchs achieve the same osmotic balance by a different mechanism; the urea compounds accumulate in the blood until there is no osmotic difference with seawater

How Terrestrial Animals Maintain Salt and Water Balance

How Terrestrial Animals Maintain Salt and Water Balance

• Animals lose water across respiratory and body surfaces, excretion of urine and elimination of feces. • Water is gained from water in food, drinking water and metabolic water. • Some desert arthropods can absorb water vapor from the air. • In desert rodents, metabolic water constitutes most of the animal’s water uptake. Dilution of Wastes a. The primary end product of protein breakdown is highly toxic urea. b. Fishes excrete urea across the gills and the abundance of water keeps it dilute. c. Terrestrial insects, reptiles and birds convert urea to nontoxic uric acid. d. Uric acid is insoluble and is excreted with little water loss. e. Uric acid can be stored in harmless crystalline form within an egg until hatching.

• Marine birds and turtles have a salt gland that secretes concentrated sodium chloride; these are important accessory glands to the kidneys that only produce very diluted urine.

Invertebrate Excretory Structures

Contractile Vacuole • These are small, spherical, intracellular vacuoles of protozoa, freshwater sponges, and radiate animals. • They are not truly excretory; ammonia and other nitrogenous wastes diffuse across the cell membrane. • The contractile vacuole is an organ of water balance; it expels excess water that enters by osmosis. • A network of channels populated with proton pumps probably surrounds them. • Contractile vacuoles are absent in marine forms that are isosmotic with seawater.

• tubular structure most common to maintain osmotic balance. • flame cells system (closed) or protonephridium is the simplest arrangement. • Planaria and other flatworms have a highly branched duct system to all parts of the body. • Fluid enters the system through specialized “flame cells” and passes through tubules to exit the body. • Rhythmical beating of a flagellar tuft creates a negative pressure that draws fluid into the tubes. • In the tubule, water and metabolites are recovered by reabsorption; wastes are left to be expelled.

• A metanephridium is an open system found in molluscs and annelids. • The tubule is open at both ends; fluid is swept into the tubule through a ciliated funnel-like opening. • A network of blood vessels to reclaim water and valuable solutes surrounds a metanephridium. • The basic process of urine formation in the tubule remains the same: withdraw valuable solutes and add waste solutes.

Arthropod Kidneys
Paired Antennal Glands of Crustaceans a. Located in the ventral part of the head, they are an advanced design of nephridia. b. They lack open nephrostomes. c. Hydrostatic pressure of the blood forms a protein-free filtrate in the end sac. d. In the tubular portion, certain salts are selectively reabsorbed or actively secreted. e. This crustacean system is similar to the vertebrate system in sequence of urine formation.

Malpighian Tubules • Insects and spiders use this system in conjunction with rectal glands. • The thin, elastic, blind tubules are closed and lack an arterial supply. • Salts, largely potassium, are actively secreted into portions of the tubule from the hemolymph. • Uric acid and other wastes continue out in the feces. • This is an especially efficient system for dry environments.

Vertebrate Kidney Function
Ancestry and Embryology 1. The earliest vertebrate kidney had segmentally arranged tubules similar to an invertebrate nephridium. 2. Each tubule opened into the coelom at a nephrostome; the other end led into a common archinephric duct. 3. This ancient kidney is called an archinephros; a similar segmented kidney is in embryos of hagfishes. 4. From the earliest time, the reproductive system from the same mesoderm used the nephric ducts. 5. Three developmental stages occur in the embryonic development of vertebrate kidneys. 6. The pronephros is observed in vertebrate embryos; it usually degenerates except in hagfish and a few bony fish species. 7. The mesonephros and metanephros (opithonephros) replace the pronephros and forms the adult kidney of fishes and amphibians. 8. The Metanephros a. This is found in adult amniotes. b. It is more caudally located and is much larger and compact. c. It contains a large number of nephric tubules. d. The ureter is a new duct that drains the system; the archinephric duct has shifted to sperm transport. 9. The three stages succeed each other embryologically and phylogenetically in amniotes.

Vertebrate Kidney Function
• The vertebrate kidney is the principal organ that regulates volume and composition of internal fluids. • The removal of metabolic wastes is incidental to its regulatory function. • Urine is formed in the nephron (funcitonal unit) by filtration, reabsorption and secretion

Structure of Human Kidney
• Kidneys comprise less than one percent of body weight but filter 20–25% of blood output. • Blood flows through nearly a million nephrons in each kidney. • The nephron begins in the renal corpuscle that contains a tuft of capillaries called the glomerulus. • Blood pressure in capillaries forces protein-free filtrate into a renal tubule. • Filtrate passes into a proximal convoluted tubule, the loop of Henle and the distal convoluted tubule. • Collecting ducts join to form the renal pelvis and carry urine through a ureter to the urinary bladder. • The blood from the dorsal aorta enters each kidney through the renal artery. • A renal artery branches to an afferent arteriole; they leave the renal corpuscle as efferent arterioles. • Efferent arterioles travel through extensive capillary networks around the proximal and distal convoluted tubules and the loop of Henle. • The capillary network collects to form the renal vein that returns blood to the vena cava.

Glomerular Filtration
• The glomerulus is a specialized mechanical filter. • Blood pressure drives a protein-free filtrate across capillary walls into the fluid-filled renal corpuscle. • Smaller solute particles are also carried across if they can fit through the slit pores of capillary walls. • Red blood cells and plasma proteins are too large to pass across. • The filtrate will undergo extensive modification before becoming urine. • About 180 liters (50 gallons) of filtrate form each day; most is reabsorbed and 1.2 liters of urine form.

Tubular Reabsorption
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About 60% of filtrate volume and nearly all glucose, amino acids, and vitamins are reabsorbed in the proximal convoluted tubule. Most reabsorption is by active transport; unique ion pumps retrieve sodium, calcium, potassium, etc. Water passively follows the osmotic gradient with active reabsorption of solutes. For most substances, there is an upper limit to reabsorption (transport maximum or renal threshold). Normally there is no glucose in urine because the transport maximum is well above the glucose level. A kidney filters 600 grams of sodium a day and retrieves 596 grams, excreting 4 grams. If human intake of sodium is higher than 4 grams, excess sodium may build in tissues and cause problems. The distal convoluted tubule carries out final adjustment of filtrate composition. About 85% of sodium absorbed by the proximal convoluted tubule is obligatory or set. In the distal convoluted tubule, sodium reabsorption is controlled by aldosterone. Aldosterone increases active reabsorption of sodium in distal tubules and decreases loss of sodium

Tubular Reabsorption
• Secretion of aldosterone is regulated by the enzyme renin, produced by the juxtaglomerular apparatus, a complex of cells located in the afferent arteriole at its junction with the glomerulus • Renin is released in response to low blood sodium level or low blood pressure. • Renin initiates a series of enzymes that result in production of angiotensin, a blood protein. • Angiotensin stimulates release of aldosterone, which increases sodium reabsorption in the distal tubule • Angiotensin also increases secretion of antidiuretic hormone (vasopressin) that promotes water conservation by the kidney. • Angiotensin increases blood pressure and stimulates thirst. • These actions reverse the circumstances that triggered the secretion of renin: sodium and water are conserved, and blood volume and pressure return to normal. • Selective pressure has restricted sodium levels in humans while sodium range is broad in rodents

Tubular Secretion
• The nephron also secretes materials into the filtrate, the reverse of tubular reabsorption. • Carrier proteins in tubular epithelial cells selectively transport substances from blood to tubule. • This allows a build up of urine concentrations of hydrogen and potassium ions, drugs, etc. • Most tubular secretion is at the distal convoluted tubule. • Marine fishes, reptiles and birds use tubular secretion far more than mammals. • Marine bony fishes actively secrete large amounts of magnesium and sulfate from seawater salts. • Uric acid is actively secreted by the tubular epithelium

Water Excretion
• The kidney closely regulates the osmotic pressure of the blood. • When fluid intake is high, the kidney excretes dilute urine and saves salts. • When fluid intake is low, the kidney conserves water and forms concentrated urine. • A dehydrated person can concentrate urine to approximately four times blood osmotic concentration

Urine concentration in mammals

• Mammal and bird kidneys produce concentrated urine by interaction between the loop of Henle and the collecting ducts. • This forms an osmotic gradient in the kidney. • In the cortex, the interstitial fluid is isosmotic with the blood. • Deep in the medulla, the osmotic concentration is four times greater than that of the blood. • High osmotic concentrations in the medulla are produced by an exchange of ions in the loop of Henle. • “Countercurrent” refers to the opposite directions of the loop of Henle, down the descending limb and up the ascending limb. • “Multiplication” describes the increasing osmotic concentration in the medulla from ion exchange between the two limbs.

Countercurrent Multiplication

• The descending limb of the loop of Henle is permeable to water but impermeable to solutes. • The ascending limb of the loop of Henle is impermeable to both water and solutes. • Sodium chloride is actively transported out of the thick portion of the ascending limb and into surrounding tissue. • As the interstitial area surrounding the loop becomes concentrated, water is withdrawn from the descending limb by osmosis. • Tubular fluid at the base of the loop is more concentrated and moves up the ascending limb where more sodium is pumped out. • The effect of active ion transport in the ascending limb is multiplied as more water is withdrawn from the descending limb and more concentrated fluid is presented to the ascending limb ion pump

Osmotic concentration of tissue fluid in mammalian kidney

• Because of high concentration of solutes surrounding a collecting duct, water is withdrawn from urine. • As urine becomes concentrated, urea diffuses out, adding to a high osmotic pressure in the kidney medulla. • The amount of water reabsorbed depends on the permeability of the walls of the distal convoluted tubule. • This is controlled by antidiuretic hormone (ADH, or vasopressin) released by the posterior pituitary. • ADH increases the permeability of the collecting duct and water diffuses outward. • In overhydration, the pituitary stops releasing ADH, pores in duct walls close, and more urine is excreted.

Temperature Regulation

Chemical Environment 1. Biochemical activities are sensitive to temperature because enzymes have an optimum temperature. 2. Temperature constraints on animals are due to their need to maintain biochemical stability. 3. At colder temperatures, metabolic reactions may be too slow to maintain activity and reproduction. 4. At too high temperatures, enzyme activity is impaired or destroyed. 5. Generally animals function between 0o and 40o C. 6. Animals may locate such habitats or develop means to stabilize their metabolism.

Ectothermy and Endothermy
• “Cold-blooded” and “warm-blooded” are not welldefined terms; fishes, insects, and reptiles basking in the sun may be warmer than mammals. • Many “warm-blooded” mammals hibernate and approach very cold temperatures. • “Poikilothermic” refers to a variable body temperature, and “homeothermic” refers to maintaining a constant body temperature; these terms likewise pose definition problems. • All animals produce some heat from cellular metabolism

• heat is conducted away as fast as it is produced. • Many ectotherms may behaviorally select areas of more favorable temperature, such as basking in the sun

Ectotherms exhibit behavioural thermoregulation

• able to generate enough heat to elevate their own temperature to a high and stable level. • Birds and mammals are the commonly recognized endotherms. • Some reptiles and fast-swimming fishes, and at times certain insects, may also be endotherms. • Endothermy allows birds and mammals to stabilize internal biochemical processes and nervous system functions. • Endotherms can also remain active in winter and exploit habitats denied to ectotherms

• Endotherms show physiological and behavioural thermoregulation

How Ectotherms Achieve Temperature Independence
• Some ectotherms behaviorally regulate body temperature. • Desert lizards exploit hour-to-hour changes in solar radiation. • In cool mornings, they bask with bodies flattened; in the hottest daytime, they retreat to burrows. • Such lizards hold body temperature between 36o and 39o C while the environment varies from 29o to 44o C. Metabolic Adjustments a. Within limits, most ectotherms can adjust metabolic rate to the prevailing temperature. b. Temperature compensation involves complex biochemical and cellular adjustments. c. This process allows a fish or salamander to sustain the same level of activity in warm or cold water.

How a lizard regulates its body temperature behaviorally

Temperature Regulation in Endotherms
• Most mammals have body temperatures between 36o and 38o C; most birds range from 40o to 42o C. • Much of an endotherm’s daily caloric intake goes to generate heat; it must eat more than an ectotherm. • Heat is lost by radiation, conduction and convection to a cooler environment, and by evaporation of water. • If an animal becomes too cool, it can generate heat by exercise or shivering, and decrease heat loss by increasing insulation.

Exchange of heat between a large mammal & its env’t on a warm day

Adaptations for Hot Environments
• Small desert animals are often fossorial, living in the ground, or nocturnal, active at night. • Lower temperatures and higher humidity of burrows also reduces water loss from evaporation. • Desert animals may also drink no water, but derive all water from food and produce dry feces. • The desert eland has many adaptations for desert living: glossy, pallid fur; fur insulation; fat tissue isolated on the back; and dropping body temperature at night. • When the body temperature reaches 41o C, it uses evaporative cooling by sweating and panting. • The desert camel has all of these adaptations perfected for desert living.

Figure 30.18

Adaptations for Cold Environments
a. Mammals and birds use decreased conductance and increased heat production to survive the cold. b. In winter, fur may increase in thickness by 50%. c. Countercurrent Heat Exchange 1) A well-insulated body can lose substantial heat through blood flowing along exposed limbs. 2) Arterial blood in the leg of an arctic mammal or bird passes in contact with returning cold blood. 3) The heat exchange all along the opposite vessels transfers nearly all body heat to the returning venous blood that returns to the body core. 4) Similar countercurrent exchange systems keep aquatic mammal flippers from losing body heat. 5) Footpads and hooves must be able to operate at near-freezing temperatures. d. Augmented muscular activity increases heat by exercise or shivering. e. Nonshivering thermogenesis uses increased oxidation of stores of brown fat. f. Small mammals live in the milder climate under the snow, a subnivean environment

Adaptive Hypothermia in Birds and Mammals
• • • • • • • • • • • • The endotherm must always have an energy supply to support its high metabolic rate. Small birds and mammals have an intense metabolism that is difficult to support on cold nights. Daily torpor is dropping body temperature when asleep or inactive; it prevents energy loss. Hummingbirds may drop body temperature at night when food supplies are low. Hibernation is prolonged and controlled dormancy. True hibernators prepare for winter by storing body fat. Entry into hibernation is gradual; the animal eventually cools to near ambient temperature. Respiration may drop from 200 breaths per minute to 4–5 per minute; heart rate from 150 to 5 beats per minute. Arousal from hibernation may require shivering and nonshivering thermogenesis. Bears, badgers, raccoons and opossums enter a prolonged sleep with little or no decrease in body temperature. This prolonged sleep is not true hibernation the animal can be awakened if disturbed. Some invertebrates and vertebrates enter of state of summer dormancy called estivation.