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HOMEOSTASIS: THERMOREGULATION & OSMOREGULATION CHAPTER 4.1 Outline Overview: Form and Function Hierarchical Organization of the Body Plane Homeostasis and Feedback loops Thermoregulation Osmoregulation Mammalian Kidney Overview: Diversity in Form and Function Anatomy is the study of the biological form of an organism Physiology is the study of the biological functions an organism performs The comparative study of animals reveals that form and function are closely correlated Fig. 40-1 Physical Constraints on Animal Size and Shape The ability to perform certain actions depends on an animal’s shape, size, and environment Evolutionary convergence reflects different species’ adaptations to a similar environmental challenge Physical laws impose constraints on animal size and shape Exchange with the Environment An animal’s size and shape directly affect how it exchanges energy and materials with its surroundings Exchange occurs as substances dissolved in the aqueous medium diffuse and are transported across the cells’ plasma membranes A single-celled protist living in water has a sufficient surface area of plasma membrane to service its entire volume of cytoplasm Fig. 40-3 Mouth Gastrovascular cavity Exchange Exchange Exchange 0.15 mm 1.5 mm (a) Single cell (b) Two layers of cells Multicellular organisms with a sac body plan have body walls that are only two cells thick, facilitating diffusion of materials More complex organisms have highly folded internal surfaces for exchanging materials In vertebrates, the space between cells is filled with interstitial fluid, which allows for the movement of material into and out of cells A complex body plan helps an animal in a variable environment to maintain a relatively stable internal environment Fig. 40-4 External environment CO2 Food O2 Mouth Respiratory system 0.5 cm 50 µm Animal body Lung tissue Nutrients Heart Cells Circulatory system 10 µm Interstitial fluid Digestive system Excretory system Lining of small intestine Kidney tubules Anus Unabsorbed matter (feces) Metabolic waste products (nitrogenous waste) Fig. 40-4a External environment CO2 O Food 2 Mouth Animal body Respiratory system Nutrients Heart Cells Circulatory system Interstitial fluid Digestive system Excretory system Anus Unabsorbed matter (feces) Metabolic waste products (nitrogenous waste) Hierarchical Organization of Body Plans Most animals are composed of specialized cells organized into tissues that have different functions Different tissues have different structures that are suited to their functions Tissues make up organs, which together make up organ systems Table 40-1 Feedback control loops maintain the internal environment in many animals Animals manage their internal environment by regulating or conforming to the external environment A regulator uses internal control mechanisms to moderate internal change in the face of external, environmental fluctuation A conformer allows its internal condition to vary with certain external changes Fig. 40-7 40 River otter (temperature regulator) Body temperature (°C) 30 20 Largemouth bass (temperature conformer) 10 0 10 20 30 Ambient (environmental) temperature (ºC) 40 Homeostasis Organisms use homeostasis to maintain a “steady state” or internal balance regardless of external environment In humans, body temperature, blood pH, and glucose concentration are each maintained at a constant level of equilibrium. Mechanisms of Homeostasis Mechanisms of homeostasis moderate changes in the internal environment For a given variable, fluctuations above or below a set point serve as a stimulus; these are detected by a sensor and trigger a response The response returns the variable to the set point Fig. 40-8 Response: Heater turned off Room temperature decreases Stimulus: Control center (thermostat) reads too hot Set point: 20ºC Stimulus: Control center (thermostat) reads too cold Room temperature increases Response: Heater turned on Homeostasis: Organ Systems The organ systems of the human body contribute to homeostasis The digestive system The respiratory system Adds oxygen to the blood Removes carbon dioxide The liver and the kidneys Takes in and digests food Provides nutrient molecules that replace used nutrients Store excess glucose as glycogen Later, glycogen is broken down to replace the glucose used The hormone insulin regulates glycogen storage The kidneys Under hormonal control as they excrete wastes and salts Feedback Loops in Homeostasis The dynamic equilibrium of homeostasis is maintained by negative feedback, which helps to return a variable to either a normal range or a set point Most homeostatic control systems function by negative feedback, where buildup of the end product shuts the system off Positive feedback loops occur in animals, but do not usually contribute to homeostasis Set points and normal ranges can change with age or show cyclic variation Homeostasis can adjust to changes in external environment, a process called acclimatization Negative Feedback Homeostatic Control Partially controlled by hormones Ultimately controlled by the nervous system Negative Feedback is the primary homeostatic mechanism that keeps a variable close to a set value Sensor detects change in environment Regulatory Center activates an effector Effector reverses the changes Positive Feedback During positive feedback, an event increases the likelihood of another event Childbirth process Urge to urinate Positive Feedback Does not result in equilibrium Does not occur as often as negative feedback Positive Feedback Copyright © The McGraw-Hill Companies, Inc. Permission required for reproduction or display. 2. Signals cause pituitary gland to release the hormone oxytocin. As the level of oxytocin increases, so do uterine contractions until birth occurs. pituitary gland + + 1. Due to uterine contractions, baby’s head presses on cervix, and signals are sent to brain. uterus Homeostasis: Bioenergentics Bioenergetics is the overall flow and transformation of energy in an animal It determines how much food an animal needs and relates to an animal’s size, activity, and environment Animals harvest chemical energy from food Energy-containing molecules from food are usually used to make ATP, which powers cellular work After the needs of staying alive are met, remaining food molecules can be used in biosynthesis Biosynthesis includes body growth and repair, synthesis of storage material such as fat, and production of gametes Homeostasis: Thermoregulation Thermoregulation is the process by which animals maintain an internal temperature within a tolerable range Endothermic animals generate heat by metabolism; birds and mammals are endotherms Ectothermic animals gain heat from external sources; ectotherms include most invertebrates, fishes, amphibians, and non-avian reptiles In general, ectotherms tolerate greater variation in internal temperature, while endotherms are active at a greater range of external temperatures Endothermy is more energetically expensive than ectothermy Fig. 40-9 (a) A walrus, an endotherm (b) A lizard, an ectotherm Homeostasis: Thermoregulation Radiation Convection Evaporation Conduction Thermoregulatory General Adaptations Insulation (major) mammals and birds Skin, feathers, fur, and blubber reduce heat flow between an animal and its environment Circulatory adaptations Regulation of blood flow near the body surface significantly affects thermoregulation Many endotherms and some ectotherms can alter the amount of blood flowing between the body core and the skin In vasodilation, blood flow in the skin increases, facilitating heat loss In vasoconstriction, blood flow in the skin decreases, lowering heat loss countercurrent gas exhangers like geese and bottleneck dolpins Thermoregulatory General Adaptations Cooling by evaporative heat loss Panting increases the cooling effect in birds and many mammals Sweating or bathing moistens the skin, helping to cool an animal down Behavioral responses Both endotherms and ectotherms use behavioral responses to control body temperature Some terrestrial invertebrates have postures that minimize or maximize absorption of solar heat Adjusting metabolic heat production Some animals can regulate body temperature by adjusting their rate of metabolic heat production Heat production is increased by muscle activity such as moving or shivering Some ectotherms can also shiver to increase body temperature Homeostasis: Mammalian Thermoregulation Hair Epidermis Sweat pore Muscle Dermis Nerve Sweat gland Hypodermis Adipose tissue Blood vessels Oil gland Regulation of Body Temperature Homeostasis: Osmoregulation Physiological systems of animals operate in a fluid environment Relative concentrations of water and solutes must be maintained within fairly narrow limits Osmoregulation regulates solute concentrations and balances the gain and loss of water Freshwater animals show adaptations that reduce water uptake and conserve solutes Desert and marine animals face desiccating environments that can quickly deplete body water Excretion gets rid of nitrogenous metabolites and other waste products Fig. 44-1 Homeostasis: Osmoregulation Osmoregulation is based largely on controlled movement of solutes between internal fluids and the external environment Cells require a balance between osmotic gain and loss of water Osmolarity, the solute concentration of a solution, determines the movement of water across a selectively permeable membrane Fig. 44-2 Selectively permeable membrane Solutes Net water flow Water Hyperosmotic side Hypoosmotic side Osmotic Challenges Marine and land animals manage differently Osmoconformers, consisting only of some marine animals, are isoosmotic with their surroundings and do not regulate their osmolarity Osmoregulators expend energy to control water uptake and loss in a hyperosmotic or hypoosmotic environment Osmoregulators gradients must expend energy to maintain osmotic Transport Epithelia in Osmoregulation Animals regulate the composition of body fluid that bathes their cells Transport epithelia are specialized epithelial cells that regulate solute movement (water-salt balance) They are essential components of osmotic regulation and metabolic waste disposal They are arranged in complex tubular networks Nitrogenous Waste Products The type and quantity of an animal’s waste products may greatly affect its water balance Among the most important wastes are nitrogenous breakdown products of proteins and nucleic acids into ammonia (NH3) Some animals convert toxic ammonia (NH3) to less toxic compounds prior to excretion Fig. 44-9 Proteins Nucleic acids Amino acids Nitrogenous bases Amino groups Most aquatic animals, including most bony fishes Ammonia Mammals, most amphibians, sharks, some bony fishes Urea Many reptiles (including birds), insects, land snails Uric acid Nitrogenous Waste Products Ammonia: Animals that excrete need lots of water They release ammonia across the whole body surface or through gills Urea The liver of mammals and most adult amphibians converts ammonia to less toxic urea Excreted via circulatory system to kidney Energetically expensive but requires less water than ammonia Nitrogenous Waste Products Uric Acid Insects, land snails, and many reptiles, including birds, mainly excrete uric acid Largely insoluble in water and can be secreted as a paste with little water loss Uric acid is more energetically expensive to produce than urea The amount of nitrogenous waste is coupled to the animal’s energy budget Type of waste depends on the evolutionary history and habitat Osmoregulation: Excretory System Excretory systems regulate solute movement between internal fluids and the external environment Most excretory systems produce urine by refining a filtrate derived from body fluids Systems that perform basic excretory functions vary widely among animal groups They usually involve a complex network of tubules Fig. 44-10 Filtration Capillary Excretory tubule Reabsorption Secretion Urine Excretion Mammalian Osmoregulation: Kidney Kidneys, the excretory organs of vertebrates, function in both excretion and osmoregulation The mammalian excretory system centers on paired kidneys, which are also the principal site of water balance and salt regulation The mammalian kidney conserves water by producing urine that is much more concentrated than body fluids Fig. 44-14a Posterior vena cava Renal artery and vein Aorta Ureter Urinary bladder Urethra (a) Excretory organs and major associated blood vessels Kidney Fig. 44-14ab Renal medulla Posterior vena cava Renal artery and vein Aorta Renal cortex Kidney Renal pelvis Ureter Urinary bladder Urethra (a) Excretory organs and major associated blood vessels Ureter (b) Kidney structure Section of kidney from a rat 4 mm Fig. 44-14b Renal medulla Renal cortex Renal pelvis Ureter (b) Kidney structure Section of kidney from a rat 4 mm Fig. 44-14cd Juxtamedullary nephron Cortical nephron 10 µm Afferent arteriole from renal artery SEM Glomerulus Bowman’s capsule Proximal tubule Peritubular capillaries Renal cortex Efferent arteriole from glomerulus Collecting duct Renal medulla To renal pelvis (c) Nephron types Branch of renal vein Loop of Henle Distal tubule Collecting duct Descending limb Ascending limb (d) Filtrate and blood flow Vasa recta Fig. 44-14e 10 µm SEM Nephrons Each kidney composed of many tubular nephrons Each nephron composed of several parts Glomerular capsule Glomerulus Proximal convoluted tubule Loop of the nephron Distal convoluted tube Collecting duct Urine production requires three distinct processes: Glomerular filtration in glomerular capsule Tubular reabsorption at the proximal convoluted tubule Tubular secretion at the distal convoluted tubule Stepwise Processing of the Blood STEP 1: Ultrafiltration in the glomerulus Filtration occurs as blood pressure forces fluid from the blood in the glomerulus into the lumen of Bowman’s capsule Filtration of small molecules is nonselective The filtrate contains salts, glucose, amino acids, vitamins, nitrogenous wastes, and other small molecules Fig. 44-14d 10 µm Afferent arteriole from renal artery SEM Glomerulus Bowman’s capsule Proximal tubule Peritubular capillaries Efferent arteriole from glomerulus Branch of renal vein Loop of Henle (d) Filtrate and blood flow Distal tubule Collecting duct Descending limb Ascending limb Vasa recta Fig. 44-15 Proximal tubule NaCl HCO3 – Nutrients H2O K+ H+ NH3 Distal tubule H2O NaCl K+ HCO3– H+ Filtrate CORTEX Loop of Henle NaCl OUTER MEDULLA H2O NaCl Collecting duct Key Active transport Passive transport Urea NaCl INNER MEDULLA H2O Stepwise Processing of the Blood STEP 2: Proximal Tubule Reabsorption of ions, water, and nutrients takes place in the proximal tubule Molecules are transported actively and passively from the filtrate into the interstitial fluid and then capillaries Some toxic materials are secreted into the filtrate The filtrate volume decreases Fig. 44-15 Proximal tubule NaCl HCO3 – Nutrients H2O K+ H+ NH3 Distal tubule H2O NaCl K+ HCO3– H+ Filtrate CORTEX Loop of Henle NaCl OUTER MEDULLA H2O NaCl Collecting duct Key Active transport Passive transport Urea NaCl INNER MEDULLA H2O Stepwise Processing of the Blood STEP 3: Descending Limb of the Loop of Henle Reabsorption of water continues through channels formed by aquaporin proteins Movement is driven by the high osmolarity of the interstitial fluid, which is hyperosmotic to the filtrate The filtrate becomes increasingly concentrated STEP 4: Ascending Limb of the Loop of Henle In the ascending limb of the loop of Henle, salt but not water is able to diffuse from the tubule into the interstitial fluid The filtrate becomes increasingly dilute Fig. 44-15 Proximal tubule NaCl HCO3 – Nutrients H2O K+ H+ NH3 Distal tubule H2O NaCl K+ HCO3– H+ Filtrate CORTEX Loop of Henle NaCl OUTER MEDULLA H2O NaCl Collecting duct Key Active transport Passive transport Urea NaCl INNER MEDULLA H2O Stepwise Processing of the Blood STEP 5: Distal Tubule The distal tubule regulates the K+ and NaCl concentrations of body fluids The controlled movement of ions contributes to pH regulation STEP 6: Collecting Duct The collecting duct carries filtrate through the medulla to the renal pelvis Water is lost as well as some salt and urea, and the filtrate becomes more concentrated Urine is hyperosmotic to body fluids Solute Gradients and Water Conservation Urine is much more concentrated than blood The cooperative action and precise arrangement of the loops of Henle and collecting ducts are largely responsible for the osmotic gradient that concentrates the urine NaCl and urea contribute to the osmolarity of the interstitial fluid, which causes reabsorption of water in the kidney and concentrates the urine The Two-Solute Model In the proximal tubule, filtrate volume decreases, but its osmolarity remains the same The countercurrent multiplier system involving the loop of Henle maintains a high salt concentration in the kidney This system allows the vasa recta to supply the kidney with nutrients, without interfering with the osmolarity gradient Considerable energy is expended to maintain the osmotic gradient between the medulla and cortex The Two-Solute Model The collecting duct conducts filtrate through the osmolarity gradient, and more water exits the filtrate by osmosis Urea diffuses out of the collecting duct as it traverses the inner medulla Urea and NaCl form the osmotic gradient that enables the kidney to produce urine that is hyperosmotic to the blood Fig. 44-16-1 Osmolarity of interstitial fluid (mOsm/L) 300 300 300 300 CORTEX H2O H2O 400 400 H2O OUTER MEDULLA H2O 600 600 900 900 H2O H2O Key Active transport Passive transport INNER MEDULLA H2O 1,200 1,200 Fig. 44-16-2 Osmolarity of interstitial fluid (mOsm/L) 300 300 100 300 100 CORTEX H2O H2O NaCl 400 H2O OUTER MEDULLA H2O Key Active transport Passive transport INNER MEDULLA H2O NaCl 200 400 NaCl NaCl 600 400 600 700 900 NaCl H2O H2O 300 900 NaCl NaCl 1,200 1,200 Fig. 44-16-3 Osmolarity of interstitial fluid (mOsm/L) 300 300 100 300 100 CORTEX H2O H2O NaCl 300 400 400 H2O NaCl 400 300 200 H2O NaCl H2O H2O NaCl NaCl OUTER MEDULLA H2O NaCl 600 400 600 600 H2O NaCl H2O H2O Urea H2O Key Active transport Passive transport INNER MEDULLA H2O 900 NaCl NaCl 700 H2O 900 Urea H2O Urea 1,200 1,200 1,200 Urine Formation and Homeostasis Excretion of hypertonic urine Dependent upon the reabsorption of water Absorbed from Loop of the nephron, and The collecting duct Osmotic gradient within the renal medulla causes water to leave the descending limb along its entire length Adaptations of the Mammalian Kidney to Diverse Environments The form and function of nephrons in various vertebrate classes are related to requirements for osmoregulation in the animal’s habitat The juxtamedullary nephron contributes to water conservation in terrestrial animals Mammals that inhabit dry environments have long loops of Henle, while those in fresh water have relatively short loops Osmoregulation and Homeostasis Mammals control the volume and osmolarity of urine The kidneys of the South American vampire bat can produce either very dilute or very concentrated urine This allows the bats to reduce their body weight rapidly or digest large amounts of protein while conserving water Fig. 44-18 Maintenance of pH and Osmolality More than 99% of sodium filtered at glomerulus is returned to blood at the distal convoluted tubule Reabsorption of sodium regulated by hormones Aldosterone Renin Atrial Natriuretic Hormone (ANH) pH adjusted by either The reabsorption of the bicarbonate ions, or The secretion of hydrogen ions 69 Hormonal Regulation: Antidiuretic Hormone The osmolarity of the urine is regulated by nervous and hormonal control of water and salt reabsorption in the kidneys Antidiuretic hormone (ADH) increases water reabsorption in the distal tubules and collecting ducts of the kidney An increase in osmolarity triggers the release of ADH, which helps to conserve water Fig. 44-19a-1 Thirst Osmoreceptors in hypothalamus trigger release of ADH. Hypothalamus ADH Pituitary gland STIMULUS: Increase in blood osmolarity Homeostasis: Blood osmolarity (300 mOsm/L) (a) Fig. 44-19a-2 Osmoreceptors in hypothalamus trigger release of ADH. Thirst Hypothalamus Drinking reduces blood osmolarity to set point. ADH Increased permeability Pituitary gland Distal tubule H2O reabsorption helps prevent further osmolarity increase. STIMULUS: Increase in blood osmolarity Collecting duct Homeostasis: Blood osmolarity (300 mOsm/L) (a) Fig. 44-19b COLLECTING DUCT LUMEN INTERSTITIAL FLUID COLLECTING DUCT CELL ADH cAMP Second messenger signaling molecule Storage vesicle Exocytosis Aquaporin water channels H2O H2O (b) ADH receptor Fig. 44-19 Osmoreceptors in hypothalamus trigger release of ADH. Thirst INTERSTITIAL FLUID COLLECTING DUCT LUMEN Hypothalamus COLLECTING DUCT CELL ADH cAMP Drinking reduces blood osmolarity to set point. ADH Increased permeability Second messenger signaling molecule Pituitary gland Storage vesicle Distal tubule Exocytosis Aquaporin water channels H2O H2O reabsorption helps prevent further osmolarity increase. Collecting duct Homeostasis: Blood osmolarity (300 mOsm/L) (a) H2O STIMULUS: Increase in blood osmolarity (b) ADH receptor Mutation in ADH production causes severe dehydration and results in diabetes insipidus Alcohol is a diuretic as it inhibits the release of ADH The Renin-Angiotensin-Aldosterone System The renin-angiotensin-aldosterone system (RAAS) is part of a complex feedback circuit that functions in homeostasis Fig. 44-21-1 Distal tubule Renin Juxtaglomerular apparatus (JGA) STIMULUS: Low blood volume or blood pressure Homeostasis: Blood pressure, volume Fig. 44-21-2 Liver Angiotensinogen Distal tubule Renin Angiotensin I ACE Juxtaglomerular apparatus (JGA) Angiotensin II STIMULUS: Low blood volume or blood pressure Homeostasis: Blood pressure, volume Fig. 44-21-3 Liver Distal tubule Angiotensinogen Renin Angiotensin I ACE Juxtaglomerular apparatus (JGA) Angiotensin II STIMULUS: Low blood volume or blood pressure Adrenal gland Aldosterone Increased Na+ and H2O reabsorption in distal tubules Arteriole constriction Homeostasis: Blood pressure, volume You should now be able to: 1. 2. 3. 4. 5. Define homeostasis and distinguish between positive and negative feedback loops Define bioenergetic Define thermoregulation and explain how endotherms and ectotherms manage their heat budgets Distinguish between the following terms: isoosmotic, hyperosmotic, and hypoosmotic; osmoregulators and osmoconformers; stenohaline and euryhaline animals Define osmoregulation, excretion 5. 6. 7. 8. 9. Compare the osmoregulatory challenges of freshwater and marine animals Describe some of the factors that affect the energetic cost of osmoregulation Using a diagram, identify and describe the function of each region of the nephron Explain how the loop of Henle enhances water conservation Describe the nervous and hormonal controls involved in the regulation of kidney function The organism in which these measurements were made is an osmo___ and a thermal___. a. b. c. d. conformer; regulator regulator; conformer conformer; conformer regulator; regulator Which is the best interpretation of the data in this graph? a. b. c. Maia, the spider crab, is an osmoconformer in saltwater but is capable of osmoregulation in freshwater. Nereis, the clam worm, is an osmoconformer in freshwater and is capable of osmoregulation in brackish water. Carcinus, the shore crab, is capable of osmoregulation in brackish water and freshwater. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings. Which of the following is an example of a negative feedback response? a. b. c. d. As the uterus contracts, more oxytocin is released to intensify uterine contractions. Meerkats bask in the sun at the beginning of the day but avoid it during the heat of the day. Sexual stimulation leads to arousal and climax. A nursing baby stimulates the release of oxytocin, which causes letdown of milk. You are a biologist studying desert animals on a day when the temperature is 110°F. You take the following body temperature measurements: snake found under a rock, 87°F; mouse in a burrow, 100°F; lizard on a rock ledge, 105°F; beetle in the leaves of a bush, 102°F. Which is most likely endothermic? a. b. c. d. Snake Mouse Lizard beetle Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings. The sea star Porcellanaster ceruleus is found exclusively in the deep sea where the water temperature is around 4°C year round. How would you classify this organism? a. b. c. d. endothermic homeotherm endothermic poikilotherm ectothermic homeotherm ectothermic poikilotherm Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings. The naked mole rat, Heterocephalus glaber, is a mammal that inhabits burrows with a stable temperature of 28 to 32°C and has the following characteristics: no fur, a poorly developed subcutaneous fat layer, no sweat glands, and skin that is highly permeable to water. Its body temperature stays only slightly above ambient (0.5°C) over a range of 12 to 37°C. How would you classify this mammal? a. b. c. d. endothermic homeotherm ectothermic poikilotherm ectothermic homeotherm endothermic poikilotherm Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings. Which of these describes urea molecules, the primary nitrogenous waste of mammals? a. b. c. d. less toxic than ammonia more soluble in water than ammonia produced mainly in cells of the kidney require less energy to produce than ammonia Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings. Kidney function requires a great deal of ATP. Transport epithelium in the nephron contains pumps for active transport of all of the following substances except a. b. c. d. e. urea. sodium ions Na+. potassium ions K+. bicarbonate ions HCO3-. hydrogen ions H+. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings. Which of the following is part of the two-solute model explaining urine production in the nephron? a. b. c. d. e. NaCl moves out of the nephron into interstitial fluid in the descending loop of Henle. Fluid entering the distal convoluted tubule is more concentrated than fluid entering the proximal convoluted tubule. Transport epithelium in the ascending loop of Henle is impermeable to water. All transport of urea is in the direction of interstitial fluid into tubule fluid. The ratio of NaCl to urea in interstitial fluid is about the same all along the length of the nephron. Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings. Which one of the following shortterm physiological phenomena (not structural adaptations) tends to lead to a decrease in the volume of urine produced? a. b. c. d. e. increase in blood pressure increase in filtration rate into the Bowman’s capsule increase in density of aquaporin channels in collecting duct decrease in blood osmolarity decrease in sodium ion reabsorption in collecting duct Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings.