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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings About this Chapter Fluid and electrolyte homeostasis Water balance Sodium balance and ECF volume Potassium balance Behavioral mechanism in salt and water balance Integrated control of volume and osmolarity Acid-base balance
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Fluid and Electrolyte Homeostasis Na + and water: ECF volume and osmolarity K + : cardiac and muscle function Ca 2+ : exocytosis, muscle contractions, and other functions H + and HCO 3 – : pH balance Body must maintain mass balance Excretion routes: kidney and lungs
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Fluid and Electrolyte Homeostasis The body’s integrated response to changes in blood volume and blood pressure Figure 20-1a
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-1b Fluid and Electrolyte Homeostasis
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-2 Water Balance in the Body
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-3 Water Balance A model of the role of the kidneys in water balance
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-4 Urine Concentration Osmolarity changes as filtrate flows through the nephron
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-5a Water Reabsorption Water movement in the collecting duct in the presence and absence of vasopressin
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-5b Water Reabsorption
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-6 Water Reabsorption The mechanism of vasopressin action Collecting duct lumen Filtrate 300 mOsm H2OH2O Exocytosis of vesicles Cross-section of kidney tubule Collecting duct cell Second messenger signal H2OH2O cAMP Storage vesicles Aquaporin-2 water pores 600 mOsM H2OH2O Medullary interstitial fluid Vasopressin receptor 600 mOsM Vasa recta H2OH2O 700 mOsM Vasopressin binds to mem- brane receptor. Receptor activates cAMP second messenger system. Cell inserts AQP2 water pores into apical membrane. Water is absorbed by osmosis into the blood. 123 4 1 2 3 4
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-6, step 1 Water Reabsorption Collecting duct lumen Filtrate 300 mOsm Cross-section of kidney tubule Collecting duct cell Medullary interstitial fluid Vasopressin receptor Vasa recta Vasopressin binds to mem- brane receptor. 1 1 600 mOsM 700 mOsM
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-6, steps 1–2 Water Reabsorption Collecting duct lumen Filtrate 300 mOsm Cross-section of kidney tubule Collecting duct cell Second messenger signal cAMP Medullary interstitial fluid Vasopressin receptor Vasa recta Vasopressin binds to mem- brane receptor. Receptor activates cAMP second messenger system. 12 1 2 600 mOsM 700 mOsM
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-6, steps 1–3 Water Reabsorption Collecting duct lumen Filtrate 300 mOsm Exocytosis of vesicles Cross-section of kidney tubule Collecting duct cell Second messenger signal cAMP Storage vesicles Aquaporin-2 water pores Medullary interstitial fluid Vasopressin receptor Vasa recta Vasopressin binds to mem- brane receptor. Receptor activates cAMP second messenger system. Cell inserts AQP2 water pores into apical membrane. 123 1 2 3 600 mOsM 700 mOsM
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-6, steps 1–4 Water Reabsorption Collecting duct lumen Filtrate 300 mOsm H2OH2O Exocytosis of vesicles Cross-section of kidney tubule Collecting duct cell Second messenger signal H2OH2O cAMP Storage vesicles Aquaporin-2 water pores 600 mOsM H2OH2O Medullary interstitial fluid Vasopressin receptor 600 mOsM Vasa recta H2OH2O 700 mOsM Vasopressin binds to mem- brane receptor. Receptor activates cAMP second messenger system. Cell inserts AQP2 water pores into apical membrane. Water is absorbed by osmosis into the blood. 123 4 1 2 3 4
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-7 Factors Affecting Vasopressin Release
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-8 Water Balance The effect of plasma osmolarity on vasopressin secretion by the posterior pituitary
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-9 Countercurrent Heat Exchanger
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-10 Water Balance Countercurrent exchange in the medulla of the kidney
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-11 Ion reabsorption Active reabsorption of ions in the thick ascending limb creates a dilute filtrate in the lumen
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Fluid and Electrolyte Balance Vasa recta removes water Close anatomical association of the loop of Henle and the vasa recta Urea increase the osmolarity of the medullary interstitium
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-12 Animation: Urinary System: Late Filtrate Processing PLAY Sodium Balance Homeostatic responses to salt ingestion
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-13 Sodium Balance Aldosterone action in principle cells Interstitial fluid Blood Aldosterone ATP Aldosterone receptor New channels P cell of distal nephron Translation and protein synthesis Proteins modulate existing channels and pumps. New pumps K+K+ Na + K+K+ K+K+ ATP K + secreted Na + reabsorbed Lumen of distal tubule Aldosterone combines with a cytoplasmic receptor. Hormone-receptor complex initiates transcription in the nucleus. New protein channels and pumps are made. Aldosterone- induced proteins modify existing proteins. Result is increased Na + reabsorption and K + secretion. 1 2 3 4 5 12 3 4 5 Transcription mRNA
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-13, step 1 Sodium Balance Interstitial fluid Blood Aldosterone receptor P cell of distal nephron Lumen of distal tubule Aldosterone combines with a cytoplasmic receptor. 1 1
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-13, steps 1–2 Sodium Balance Interstitial fluid Blood Aldosterone receptor P cell of distal nephron Lumen of distal tubule Aldosterone combines with a cytoplasmic receptor. Hormone-receptor complex initiates transcription in the nucleus. 1 2 12 Transcription mRNA
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-13, steps 1–3 Sodium Balance Interstitial fluid Blood Aldosterone ATP Aldosterone receptor P cell of distal nephron Translation and protein synthesis Lumen of distal tubule Aldosterone combines with a cytoplasmic receptor. Hormone-receptor complex initiates transcription in the nucleus. New protein channels and pumps are made. 1 2 3 12 3 Transcription mRNA New channels New pumps
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-13, steps 1–4 Sodium Balance Interstitial fluid Blood Aldosterone ATP Aldosterone receptor P cell of distal nephron Translation and protein synthesis ATP Lumen of distal tubule Aldosterone combines with a cytoplasmic receptor. Hormone-receptor complex initiates transcription in the nucleus. New protein channels and pumps are made. Aldosterone- induced proteins modify existing proteins. 1 2 3 4 12 3 4 Transcription mRNA New channels Proteins modulate existing channels and pumps. New pumps
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-13, steps 1–5 Sodium Balance Interstitial fluid Blood Aldosterone ATP Aldosterone receptor P cell of distal nephron Translation and protein synthesis K+K+ Na + K+K+ K+K+ ATP K + secreted Na + reabsorbed Lumen of distal tubule Aldosterone combines with a cytoplasmic receptor. Hormone-receptor complex initiates transcription in the nucleus. New protein channels and pumps are made. Aldosterone- induced proteins modify existing proteins. Result is increased Na + reabsorption and K + secretion. 1 2 3 4 5 12 3 4 5 Transcription mRNA New channels Proteins modulate existing channels and pumps. New pumps
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-14 Sodium Balance The renin-angiotensin-aldosterone pathway
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-15 Sodium Balance Decreased blood pressure stimulates renin secretion
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-16 Sodium Balance Action of natriuretic peptides
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Potassium Balance Regulatory mechanisms keep plasma potassium in narrow range Aldosterone plays a critical role Hypokalemia Muscle weakness and failure of respiratory muscles and the heart Hyperkalemia Can lead to cardiac arrhythmias Causes include kidney disease, diarrhea, and diuretics
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Behavioral Mechanisms Drinking replaces fluid loss Low sodium stimulates salt appetite Avoidance behaviors help prevent dehydration Desert animals avoid the heat
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Disturbances in Volume and Osmolarity Animation: Fluid, Electrolyte, and Acid/Base Balance: Water Homeostasis PLAY Figure 20-17
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Volume and Osmolarity
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Volume and Osmolarity
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Volume and Osmolarity
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Blood volume/ Blood pressure Osmolarity Granular cells GFR Flow at macula densa accompanied by CVCC Parasympathetic output Sympathetic output Heart ForceRate Cardiac output Vasoconstriction Peripheral resistance Angiotensinogen ANG I ACE ANG II Aldosterone Na + reabsorption Distal nephron Distal nephron Vasopressin release from posterior pituitary Arterioles Volume H 2 O reabsorption H 2 O intake Thirst Volume conserved Hypothalamus Adrenal cortex DEHYDRATION CARDIOVASCULAR MECHANISMS RENIN-ANGIOTENSIN SYSTEM RENAL MECHANISMS HYPOTHALAMIC MECHANISMS Carotid and aortic baroreceptors Hypothalamic osmoreceptors Atrial volume receptors; carotid and aortic baroreceptors Osmolarity Blood pressure + + + + + + + + + + + Renin osmolarity inhibits Figure 20-18 Volume and Osmolarity Homeostatic compensation for severe dehydration
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-18 (1 of 6) Volume and Osmolarity Blood volume/ Blood pressure CVCC Parasympathetic output Sympathetic output Heart ForceRate Cardiac output Vasoconstriction Peripheral resistance Arterioles DEHYDRATION CARDIOVASCULAR MECHANISMS Carotid and aortic baroreceptors Blood pressure
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-18 (2 of 6) Volume and Osmolarity Blood volume/ Blood pressure Osmolarity accompanied by Distal nephron Vasopressin release from posterior pituitary Volume H 2 O reabsorption H 2 O intake Thirst Hypothalamus DEHYDRATION HYPOTHALAMIC MECHANISMS Hypothalamic osmoreceptors Atrial volume receptors; carotid and aortic baroreceptors Osmolarity Blood pressure + +
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-18 (3 of 6) Volume and Osmolarity Blood volume/ Blood pressure Granular cells GFR Flow at macula densa Angiotensinogen ANG I ACE ANG II Volume conserved DEHYDRATION RENIN-ANGIOTENSIN SYSTEM RENAL MECHANISMS + + Renin
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-18 (4 of 6) Volume and Osmolarity Blood volume/ Blood pressure Osmolarity Granular cells accompanied by CVCC Angiotensinogen ANG I ACE ANG II Aldosterone Na + reabsorption Distal nephron Vasopressin release from posterior pituitary Arterioles Thirst Adrenal cortex DEHYDRATION RENIN-ANGIOTENSIN SYSTEM + + + + + + Renin osmolarity inhibits
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-18 (5 of 6) Volume and Osmolarity Blood volume/ Blood pressure Osmolarity Granular cells GFR Flow at macula densa accompanied by CVCC Parasympathetic output Sympathetic output Vasoconstriction Angiotensinogen ANG I ACE ANG II Aldosterone Na + reabsorption Distal nephron Distal nephron Vasopressin release from posterior pituitary Arterioles Volume H 2 O reabsorption H 2 O intake Thirst Volume conserved Adrenal cortex DEHYDRATION RENIN-ANGIOTENSIN SYSTEM RENAL MECHANISMS Osmolarity Blood pressure + + + + + + + + + Renin osmolarity inhibits
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-18 (6 of 6) Volume and Osmolarity Blood volume/ Blood pressure Osmolarity Granular cells GFR Flow at macula densa accompanied by CVCC Parasympathetic output Sympathetic output Heart ForceRate Cardiac output Vasoconstriction Peripheral resistance Angiotensinogen ANG I ACE ANG II Aldosterone Na + reabsorption Distal nephron Distal nephron Vasopressin release from posterior pituitary Arterioles Volume H 2 O reabsorption H 2 O intake Thirst Volume conserved Hypothalamus Adrenal cortex DEHYDRATION CARDIOVASCULAR MECHANISMS RENIN-ANGIOTENSIN SYSTEM RENAL MECHANISMS HYPOTHALAMIC MECHANISMS Carotid and aortic baroreceptors Hypothalamic osmoreceptors Atrial volume receptors; carotid and aortic baroreceptors Osmolarity Blood pressure + + + + + + + + + + + Renin osmolarity inhibits
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Acid-Base Balance Normal pH of plasma is 7.38–7.42 H + concentration is closely regulated Changes can alter tertiary structure of proteins Abnormal pH affects the nervous system Acidosis: neurons become less excitable and CNS depression Alkalosis: hyperexcitable pH disturbances Associated with K + disturbances
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-19 Acid-Base Balance Hydrogen balance in the body
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Acid and Base Input Acid Organic acids Diet and intermediates Under extraordinary conditions Metabolic organic acid production can increase Ketoacids Diabetes Production of CO 2 Acid production Base Few dietary sources of bases
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings pH Homeostasis Buffers moderate changes in pH Cellular proteins, phosphate ions, and hemoglobin Ventilation Rapid 75% of disturbances Renal regulation Directly excreting or reabsorbing H + Indirectly by change in the rate at which HCO 3 – buffer is reabsorbed or excreted
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-20 pH Disturbances The reflex pathway for respiratory compensation of metabolic acidosis
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-21 pH Disturbances Overview of renal compensation for acidosis
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Renal Compensation: Transporters Apical Na + -H + antiport Basolateral Na + -HCO 3 – symport H + -ATPase H + -K + -ATPase Na + -NH 4 + antiport
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-22 Renal Compensation Proximal tubule H + secretion and the reabsorption of filtered HCO 3 – Peritubular capillary Interstitial fluid Reabsorbed Filtration Proximal tubule cell Glomerulus HCO 3 – H + + HCO 3 – HCO 3 – CO 2 + H 2 O H+H+ Na + Secreted H + Na + HCO 3 – Na + Bowman’s capsule H 2 O + CO 2 Filtered HCO 3 – + H + Secreted H + and NH 4 + will be excreted Na + KG HCO 3 – Na + NH 4 + HCO 3 – Na + Glutamine CA Na + -H + antiport secretes H +. H + in filtrate combines with filtered HCO 3 – to form CO 2. CO 2 diffuses into cell and combines with water to form H + and HCO 3 –. H + is secreted again and excreted. HCO 3 – is reabsorbed. Glutamine is metabolized to ammonium ion and HCO 3 –. NH 4 + is secreted and excreted. HCO3– is reabsorbed. 1 2 4 3 6 7 5 8 1 2 4 3 6 7 5 8
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-22, step 1 Renal Compensation Peritubular capillary Interstitial fluid Filtration Proximal tubule cell Glomerulus H+H+ Na + Secreted H + Bowman’s capsule Na + Na + -H + antiport secretes H +. 1 1
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-22, steps 1–2 Renal Compensation Peritubular capillary Interstitial fluid Filtration Proximal tubule cell Glomerulus HCO 3 – H+H+ Na + Secreted H + Bowman’s capsule H 2 O + CO 2 Filtered HCO 3 – + H + Na + Na + -H + antiport secretes H +. H + in filtrate combines with filtered HCO 3 – to form CO 2. 1 2 1 2
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-22, steps 1–3 Renal Compensation Peritubular capillary Interstitial fluid Filtration Proximal tubule cell Glomerulus HCO 3 – H + + HCO 3 – CO 2 + H 2 O H+H+ Na + Secreted H + Bowman’s capsule H 2 O + CO 2 Filtered HCO 3 – + H + Na + Na + -H + antiport secretes H +. H + in filtrate combines with filtered HCO 3 – to form CO 2. CO 2 diffuses into cell and combines with water to form H + and HCO 3 –. 1 2 3 1 2 3 CA
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-22, steps 1–4 Renal Compensation Peritubular capillary Interstitial fluid Filtration Proximal tubule cell Glomerulus HCO 3 – H + + HCO 3 – CO 2 + H 2 O H+H+ Na + Secreted H + Bowman’s capsule H 2 O + CO 2 Filtered HCO 3 – + H + Secreted H + will be excreted Na + Na + -H + antiport secretes H +. H + in filtrate combines with filtered HCO 3 – to form CO 2. CO 2 diffuses into cell and combines with water to form H + and HCO 3 –. H + is secreted again and excreted. 1 2 4 3 1 2 4 3 CA
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-22, steps 1–5 Renal Compensation Peritubular capillary Interstitial fluid Reabsorbed Filtration Proximal tubule cell Glomerulus HCO 3 – H + + HCO 3 – HCO 3 – CO 2 + H 2 O H+H+ Na + Secreted H + Na + HCO 3 – Na + Bowman’s capsule H 2 O + CO 2 Filtered HCO 3 – + H + Secreted H + will be excreted Na + Na + -H + antiport secretes H +. H + in filtrate combines with filtered HCO 3 – to form CO 2. CO 2 diffuses into cell and combines with water to form H + and HCO 3 –. H + is secreted again and excreted. HCO 3 – is reabsorbed. 1 2 4 3 5 1 2 4 3 5 CA
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-22, steps 1–6 Renal Compensation Peritubular capillary Interstitial fluid Reabsorbed Filtration Proximal tubule cell Glomerulus HCO 3 – H + + HCO 3 – HCO 3 – CO 2 + H 2 O H+H+ Na + Secreted H + Na + HCO 3 – Na + Bowman’s capsule H 2 O + CO 2 Filtered HCO 3 – + H + Secreted H + will be excreted Na + KG HCO 3 – NH 4 + Glutamine Na + -H + antiport secretes H +. H + in filtrate combines with filtered HCO 3 – to form CO 2. CO 2 diffuses into cell and combines with water to form H + and HCO 3 –. H + is secreted again and excreted. HCO 3 – is reabsorbed. Glutamine is metabolized to ammonium ion and HCO 3 –. 1 2 4 3 6 5 1 2 4 3 6 5 CA
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-22, steps 1–7 Renal Compensation Peritubular capillary Interstitial fluid Reabsorbed Filtration Proximal tubule cell Glomerulus HCO 3 – H + + HCO 3 – HCO 3 – CO 2 + H 2 O H+H+ Na + Secreted H + Na + HCO 3 – Na + Bowman’s capsule H 2 O + CO 2 Filtered HCO 3 – + H + Secreted H + and NH 4 + will be excreted Na + KG HCO 3 – NH 4 + Glutamine Na + -H + antiport secretes H +. H + in filtrate combines with filtered HCO 3 – to form CO 2. CO 2 diffuses into cell and combines with water to form H + and HCO 3 –. H + is secreted again and excreted. HCO 3 – is reabsorbed. Glutamine is metabolized to ammonium ion and HCO 3 –. NH 4 + is secreted and excreted. 1 2 4 3 6 7 5 1 2 4 3 6 7 5 CA
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-22, steps 1–8 Renal Compensation Peritubular capillary Interstitial fluid Reabsorbed Filtration Proximal tubule cell Glomerulus HCO 3 – H + + HCO 3 – HCO 3 – CO 2 + H 2 O H+H+ Na + Secreted H + Na + HCO 3 – Na + Bowman’s capsule H 2 O + CO 2 Filtered HCO 3 – + H + Secreted H + and NH 4 + will be excreted Na + KG HCO 3 – Na + NH 4 + HCO 3 – Na + Glutamine Na + -H + antiport secretes H +. H + in filtrate combines with filtered HCO 3 – to form CO 2. CO 2 diffuses into cell and combines with water to form H + and HCO 3 –. H + is secreted again and excreted. HCO 3 – is reabsorbed. Glutamine is metabolized to ammonium ion and HCO 3 –. NH 4 + is secreted and excreted. HCO3– is reabsorbed. 1 2 4 3 6 7 5 8 1 2 4 3 6 7 5 8 CA
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Figure 20-23 Intercalated Cells Role of intercalated cells in acidosis and alkalosis
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Acid-Base Balance Animation: Fluid, Electrolyte, and Acid/Base Balance: Acid/Base Homeostasis PLAY
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Summary Fluid and electrolyte homeostasis Water balance Vasopressin, aquaporin, osmoreceptors, countercurrent multiplier, and vasa recta Sodium balance Aldosterone, principal cells, ANG I and II, renin, angiotensinogen, ACE, and ANP Potassium balance Hyperkalemia and hypokalemia
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Copyright © 2007 Pearson Education, Inc., publishing as Benjamin Cummings Summary Behavioral mechanisms Integrated control of volume and osmolarity Acid-base balance Buffers, ventilation, and kidney Acidosis and alkalosis Intercalated cells
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