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4/2/2015 Chapter 44: Osmoregulation & Excretion 1. Osmoregulation 2. Nitrogenous Wastes 3. Excretory Processes 4. Hormonal Control of Osmoregulation & Excretion 1. Osmoregulation Balancing Uptake & Loss of Water, Solutes Osmoregulation is the process of balancing the uptake and loss of water as well maintaining solute concentrations within acceptable levels: • the key factor in this balance is osmosis, the diffusion of water from high to low concentration across a semi-permeable membrane 1 4/2/2015 Osmosis & Osmolarity Differences in osmolarity (the total solute concentration) across a membrane permeable only to the water will result in osmosis – the diffusion of water Selectively permeable from higher to lower membrane Solutes Water concentration: The side with higher [solute] will gain water and the side with lower [solute] will lose water. Hyperosmotic side: Hypoosmotic side: Higher solute concentration • Lower free H O 2 concentration • • • Lower solute concentration Higher free H2O concentration Net water flow Osmolarity & Aquatic Animals Some aquatic animals such as marine invertebrates are osmoconformers that are isoosmotic with their surroundings and thus don’t regulate their osmolarity. However most are osmoregulators that expend energy to regulate their osmolarity. • most aquatic animals are stenohaline and cannot tolerate large fluctuations in the osmolarity of their environment • some animals such as salmon and bull sharks are euryhaline and can survive large fluctuations in the osmolarity of their environment Osmoregulation in Marine Fish Marine bony fish are hypoosmotic to sea water and thus will lose water and take in excess salt. To counter this bony fish: (a) Osmoregulation in a marine fish • ingest sea water Gain of water and salt ions from food • reduce the amount of water excreted in the urine • excrete excess salt from gills and kidneys Excretion of salt ions from gills Osmotic water loss through gills and other parts of body surface Key SALT WATER Gain of water and salt ions from drinking seawater Water Salt Excretion of salt ions and small amounts of water in scanty urine from kidneys 2 4/2/2015 Osmoregulation in Freshwater Fish Freshwater fish are hyperosmotic to sea water and thus will gain water and lose salt. To counter this freshwater fish: (b) Osmoregulation in a freshwater fish Gain of water and some ions in food Uptake of salt ions by gills Osmotic water gain through gills and other parts of body surface • excrete large amounts of water from the kidneys Key Water Salt • uptake salt in the gills FRESH WATER Excretion of salt ions and large amounts of water in dilute urine from kidneys Transport Epithelia in Osmoregulation Nostril with salt secretions • transport epithelia are cell layers specialized for moving solutes in specific directions typically arranged in tubular networks such as the nasal glands of marine birds to excrete excess salt Secretory cell of transport epithelium Vein Artery Lumen of secretory tubule Nasal salt gland Capillary Secretory tubule Salt ions Transport epithelium Blood Salt flow secretion Central duct 2. Nitrogenous Wastes 3 4/2/2015 Forms of Nitrogenous Waste All animals produce nitrogenous waste in the form of ammonia (NH3) due to the break down of proteins and nucleic acids. Ammonia is quite toxic and thus must be handled in 1 of 2 ways: 1. rapid elimination from the body 2. conversion to the less toxic nitrogen compounds urea or uric acid ammonia urea Proteins Nucleic acids • most aquatic animals can rapidly eliminate ammonia into the surrounding water Nitrogenous bases Amino acids Amino groups Most aquatic animals, including most bony fishes Ammonia Mammals, most amphibians, sharks, some bony fishes Urea uric acid Many reptiles (including birds), insects, land snails Uric acid • mammals, amphibians and some aquatic animals expend energy to convert ammonia to urea which requires more water for excretion than uric acid • reptiles, birds & insects convert ammonia to uric acid which is more energetically expensive than urea but requires little water to be excreted 3. Excretory Processes 4 4/2/2015 1 Capillary Filtration Filtrate Excretory systems get rid of nitrogenous waste and osmoregulate through a series of general steps: Excretory tubule 2 Excretory Systems in General Reabsorption FILTRATION – removal of soluble portion of body fluid to form crude filtrate REABSORPTION – reclaiming water and 3 Secretion valuable solutes Urine SECRETION – transferring additional wastes to the filtrate 4 Excretion EXCRETION – elimination of urine from body Protonephridia Protonephridia are a simple type of excretory system consisting of networks of dead-end tubules that excrete a dilute waste fluid into excretory tubules: • flame bulbs with cilia move fluid through perforated cap cells that filter out waste into excretory tubules Tubules of protonephridia INTERSTITIAL FLUID Cilia Cap cell Tubule cell Flame bulb Tubule • found in flatworms Opening in body wall Metanephridia Metanephridia are excretory organs in segmented worms (annelids), and some arthrophods and molluscs: Coelom Components of a metanephridium: Collecting tubule Internal opening Bladder External opening Capillary network • blood filtrate collects in the coelom and is transferred via cilia into a collecting tubule • reabsorption & secretion across the tubule results in a dilute urine that is released out of the body 5 4/2/2015 Malpighian Tubules Digestive tract Rectum Intestine Midgut (stomach) Salt, water, and nitrogenous wastes Hindgut Malpighian tubules To anus Feces and urine In most insects & other terrestrial arthropods, excretion occurs via Malpighian tubules: • nitrogenous & other wastes are transferred to Malpighian tubules connected to the gut Malpighian tubule Rectum Reabsorption • waste then passes out of body through the anus HEMOLYMPH Kidneys Excretory Organs • kidneys are the main excretory organs in vertebrates Posterior vena cava Renal artery and vein Kidney Kidney Structure Renal cortex Renal medulla Renal artery Renal vein Aorta Ureter Ureter Urinary bladder Renal pelvis Urethra Nephrons – the Function Unit of the Kidney Afferent arteriole from renal artery Glomerulus Bowman’s capsule Nephron Structure Cortical nephron Proximal tubule Collecting duct Branch of renal vein Renal cortex Descending limb Vasa recta Loop of Henle Ascending limb 200 µm Distal tubule Efferent arteriole from glomerulus Juxtamedullary nephron Peritubular capillaries Renal Blood vessels from a human kidney. medulla Arterioles and peritubular capillaries appear pink; glomeruli appear yellow. 6 4/2/2015 Afferent arteriole from renal artery Glomerulus & Bowman’s Capsule Glomerulus Bowman’s capsule Proximal tubule Peritubular capillaries Distal tubule Fluid from the blood leaks from the glomerulus into Bowman’s capsule: Efferent arteriole from glomerulus Branch of renal vein Descending limb Vasa recta Loop of Henle Ascending limb 200 µm Collecting duct Blood vessels from a human kidney. Arterioles and peritubular capillaries appear pink; glomeruli appear yellow. • the crude filtrate produced contains salts, glucose, amino acids, vitamins, nitrogenous wastes, & other small molecules The Proximal Tubule proximal tubule The following occurs in the proximal tubule: • reabsorption of water, minerals & nutrients (e.g., glucose) • some toxins are secreted into the tubule • these activities result in a more concentrated filtrate The Descending Loop of Henle descending limb of the Loop of Henle The following occurs in the descending limb of the Loop of Henle: • reabsorption of water by diffusion through aquaporin channels • interstitial fluid is hyperosmotic relative to filtrate • further concentrates filtrate 7 4/2/2015 The Ascending Loop of Henle ascending limb of the Loop of Henle The following occurs in the ascending limb of the Loop of Henle: • reabsorption of salt ions • interstitial fluid is hypoosmotic relative to filtrate • filtrate becomes more dilute The Distal Tubule distal tubule The following occurs in the distal tubule: • Na+, K+ & Cl- and other ions are transported in or out of the filtrate to maintain osmotic balance • H+ and HCO3- ions are transported in or out of filtrate to maintain pH balance The Collecting Duct collecting duct The following occurs in the collecting duct: • reabsorption of water and valuable solutes • resulting urine is conducted to the renal medulla on its way to the renal pelvis and ureter 8 4/2/2015 1 Proximal tubule H+ H2O NaCl K+ NH 3 NaCl H2O OUTER MEDULLA NaCl 3 Thin segment of ascending limb NaCl Active transport Passive transport 300 OUTER MEDULLA INNER MEDULLA Urea H2O mOsm/L 100 300 100 Active transport Passive transport 5 Collecting duct INNER MEDULLA 300 CORTEX H+ 3 Thick segment of ascending limb 2 Descending limb of loop of Henle H2O Salts (NaCl and others) HCO3 H+ Urea Glucose, amino acids Some drugs HCO 3 Interstitial fluid CORTEX Filtrate Summary of Nephron Function 4 Distal tubule NaCl Nutrients K+ HCO3 H2O NaCl H2O H2O 400 H2O H2O NaCl 200 NaCl 600 NaCl H2O NaCl H2O 900 NaCl H2O NaCl 1,200 300 300 400 400 H2O 400 700 H2O NaCl H2O NaCl H2O 600 600 1,200 1,200 H2O Urea H2O Urea H2O Urea 900 Osmolarity in the Cortex vs Medulla The osmolarity of the interstitial fluids surrounding the nephron increases from cortex to medulla. This gradient is responsible for the passive transport of water and solutes in various sections of the nephron. 4. Hormonal Control of Osmoregulation & Excretion 9 4/2/2015 Antidiuretic Hormone Osmoreceptors in hypothalamus trigger release of ADH. Hypothalamus Hypothalamus generates thirst. Posterior pituitary • antidirutic hormone or ADH (aka vasopressin) released by the posterior pituitary gland stimulates Drinking of water greater water reabsorption in the kidney blood when osmolarity increases due to dehydration Distal tubule ADH Blood osmolarity increases (such as after sweating profusely). Collecting duct H2O reabsorption NORMAL BLOOD OSMOLARITY (300 mOsm/L) ADH receptor LUMEN ADH cAMP …more on ADH Collecting duct COLLECTING DUCT CELL • the effect of ADH is to increase the number of aquaporin water channels on the apical surface of collecting duct epithelial cells Second messenger Protein kinase A Storage vesicle Exocytosis Aquaporin water channel • this results in more water reabsorption from the collecting duct to the interstitial fluid H2O H2O Renin-Angiotensin-Aldosterone System (RAAS) and H O are 5 Nareabsorbed. + • the RAAS is a complex hormonal feedback system that increases water reabsorption in the kidney in response decreased blood pressure Atrial natriuretic peptide (ANP) inhibits renin release when blood pressure rises. 4 2 Aldosterone Adrenal gland NORMAL BLOOD PRESSURE AND VOLUME Blood pressure or blood volume drops (for example, due to dehydration or blood loss). Arterioles constrict. 3 Angiotensin II ACE Angiotensin I 2 Distal tubule JGA releases renin. 1 Sensors in JGA detect decrease. Renin Angiotensinogen Liver Juxtaglomerular apparatus (JGA) 10