Lecture 23, 07 Nov 2006 Vertebrate Physiology ECOL 437 (MCB/VetSci 437) Univ. of Arizona, Fall 2006 Housekeeping, 07 Nov 2006 Kidney/Urine Lab on 08 November, Term Paper drafts (2 with codenames) Don’t forget to do seminar write-up! (due 10 November) (Eckert 14-17) Upcoming Readings Kevin Bonine & Kevin Oh today: Text, Ch. 25-27 (osmoregulation, kidney) Wed 08 Nov: Kidney/Urine lab, (Heart), Term Paper Thurs 09 Nov: Text, chapter 25-28, kidney, excretion, deserts Tues 14 Nov: begin 4+5 (feeding, nutrition, metabolism)? Thurs 16 Nov: Exam 3 1. Osmoregulation (Chap 25-26) 2. Kidney Function (Chap 27) 1 Lab oral presentations 08 Nov 9am – none 2pm – none 2 http://eebweb.arizona.edu/eeb_course_websites.htm Vertebrate Physiology 437 Osmoregulation -Ionic and Osmotic Balance Osmoregulation -Kidney Function (Eckert, 14-18) 3 Osmoregulation 4 Obligatory Osmotic Exchanges -life arose in salty sea 1-Gradients -Frog in freshwater -Fish in ocean -extracellular fluids ~ similar 2-Surface-to-Volume Ratio -Small animals dehydrate or hydrate more rapidly -dist’n limited by temperature and osmotic pressure (dehydration, ionic composition) -Skin, and Respiratory surface (higher metabolism with higher per/gram respiratory surface) -terrestrial organisms (and their descendents) regulate internal environment (homeostasis) 3-Integument Permeability -Transcellular or Paracellular -Aquaporins = water channel proteins -salt and water regulation (waste excretion) -kidneys, salt glands, gills 5 -Frogs vs. Lizards, Pelvic Patch etc. 6 1 Obligatory Osmotic Exchanges Osmoregulation 4-Feeding, Metabolism, Excretion -metabolic waste products ammonia, urea, etc. -metabolic water (desert!) -ingestion of salts -kidneys, salt glands, gills (more later) -Water Breathing 1. Fresh Ambystoma tigrinum Blood osmolarity 200-300 mosm/L Water ~ 50 mosm/L 5-Respiration -internalize respiratory surface -temporal countercurrent system (dry and cool IN, becomes moist and warm; recover) (countercurrent blood flow also) -temperature regulation vs. water conservation -ectotherm vs. endotherm (in deserts) - hyperosmotic animals, danger of swelling, losing salts get their water across skin dilute urine active uptake of salts across epithelium fish gills, frog skin, etc. 7 Osmoregulation 8 Osmoregulation -Water Breathing -Air Breathing 2. Salt (~1,000 mosm/L) Have to lose water to allow gas exchange - Marine reptiles and marine birds can drink seawater and secrete salts in high [ ] Most marine vertebrates hypo-osmotic (e.g., teleost or bony fishes) - SALT GLANDS - - Mammals rely on kidney danger of losing water, gaining too many salts drink saltwater excess salts actively secreted (gills, kidneys) chloride cells for salt secretion (Pelis et al. paper) (14-8) 9 10 Hill et al. 2004, Fig 26.15 (16-7) Mouse-to-Elephant Curve 4g shrew eats 2g/day elephant is 1 million x larger 11 12 2 Camel Water K-rat Free Preformed Oryx Metabolic C6H12O6 + 6O2 ÅÆ 6CO2 + 6H20 Hopping Mouse 13 14 ORGANS THAT CONTRIBUTE TO OSMOREGULATION IN VERTEBRATES K-rat Lab rats Group Osmoregulatory Organs Fish Kidneys Gills Bladder Intestine Kidneys Gills Bladder Skin Intestine Amphibians Reptiles Kidneys Salt Glands Intestine Birds Kidneys Salt Glands Intestines Mammals Kidneys 15 Osmoregulation 16 Water -Air Breathing Desert Mammals Behavior and Physiology (Eckert 14-9) Lose water: evaporation urine feces salt glands Role of microhabitat eyes Eleutherodactylus coqui Pough et al., 2001 Alter behavior and physiology to minimize water loss Water balance limits activity in time and space Kangaroo Rat -Reduce Activity -Remain in Cool Burrow -Highly concentrated urine -Very dry feces (rectal absorption) -Metabolic water 17 Amphibs lose most water via evaporation - cutaneous resistance 1 dried mucus 2 cocoon 3 wax 18 3 Water Phyllomedusa Phyllomedusa sauvagi Chuckwalla Less evap. (lizards have more lipids in skin) Monkey Tree Frog Anolis lizard Alligator Softshell Turtle More evap. Pough et al., 2001 Bufo, Spadefoots, Rana 19 (free water surface) Pough et al., 2001 20 Osmoregulatory Mechanisms Apical surface (faces lumen and outside world) Basal surface (faces body and extracellular fluid) Kidney Function - Active movement of ions/salts requires ATP - Movement of water follows movement of ions/salts Volume Regulation Osmotic Regulation Ionic Regulation How do these differ? 21 Gradients established and used…to move ions, water 22 (14-11) Fish Gills active Chloride cells involved in osmoregulation -(recall lab paper on smolting) -lots of mitochondria to power ATPases -mechanism similar in nasal glands (birds and reptiles), and shark rectal gland (14-14) passive 2 1 4 (Eckert 14-12) Mammalian Kidney 3 5 23 24 4 (Eckert 14-17) (Eckert 14-17) Mammalian Kidney -Paired -1% body mass -20% blood flow Kidney Functions: -Osmoregulation -Blood volume regulation -Maintain proper ion concentrations -Dispose of metabolic waste products -pH regulation (at ~ 7.4) -Dispose of toxins and foreign substances -urine contains: water metabolic byproducts (e.g., urea) excess salts etc. -from ureter to urinary bladder (smooth muscle, sphincter, inhibition) -out via urethra during micturition How does the kidney accomplish this? 25 26 Mammalian Kidney Anatomy FUNCTIONAL UNIT (~ 1 million) extracellular intracellular (Eckert 14-17) Urine 27 28 Hill et al. 2004, Fig. 27.6 Nephron Anatomy 1 -Proximal tubule 2 -Loop of Henle -descending -ascending 3 -Distal tubule -numerous nephrons empty into collecting duct -collecting ducts empty into renal pelvis 29 (Eckert 14-18) 30 5 Nephron Anatomy 1 2 6 3 7 4 5 Vasa recta Countercurrent exchange 31 Knut Schmidt_Nielsen 1997 32 Knut Schmidt_Nielsen 1997 Kidney Processes- overview Mosm = x1000 U/P Filtration plus secretion 1. FILTRATION blood --> filtrate 2. REABSORPTION filtrate --> blood + 3. SECRETION blood --> filtrate All 3 involved in final Urine Composition Dipodomys (Eckert 14-21) 34 Knut Schmidt_Nielsen 1997 33 Humans: 125 ml/min or 180 L/day (14-20) Hill et al. 2004, Fig 25.7 35 Sympathetic innervation tends to constrict 36 6 Filtration: Bowman’s capsule Bowman’s capsule 3 layers 1. Glomerular endothelial cells -100x leakier than other capillary walls 2. Basement membrane -negatively charged glycoproteins -repel plasma proteins by charge 3. Epithelial cells -podocytes create slits (Eckert 14-23) Filtrate = protein-free and cell-free plasma Glomerular Filtration Rate (GFR) Humans: 125 ml/min or 180 L/day (60x plasma vol.) 37 38 Bowman’s capsule proteins and larger molecules remain About 20% of the plasma and solutes that enter glomerulus end up in BC (Eckert 14-22) 39 7