31st Lecture Fri 03 April 2009 Housekeeping, Wed 01 April 2009 Readings Today, Mon 30 Mar: Ch 26 (Ionic and Osmotic Balance ) Wed 01 April: Ch 27 (Osmoregulation) LAB: Circulation; Lillywhite Fri 03 April: Ch 27 (Osmoregulation) Vertebrate Physiology ECOL 437 (MCB/VetSci 437) Univ. of Arizona, spring 2009 Kevin Bonine & Kevin Oh Osmoregulation Fri 13 Feb = Exam 1 Chapter 26 & 27 1 Research Proposal Meetings Lab discussion leaders: 01 Apr Lab discussion leaders: 08 Apr 1pm – Fred, Jarrett 1pm – xx 3pm – ?? 3pm – xx 2 Vertebrate Galapagos Marine Ecology (ECOL 496O/596O) Osmoregulation Summer Session II: July 7-Aug 1, 2009 •Spend one month this summer in the Galapagos Islands, Ecuador! •Visit seven of the most spectacular islands in the archipelago •Do a service project with children at a local school and the Galapagos National Park •Do a field ecology project and learn about Galapagos ecology and evolution •Earn 3-6 units of graduate or undergraduate credit For more information: http://www.eebweb.arizona.edu/courses/galapagos/ Katrina Mangin, mangin@email.arizona.edu, 520-626-5076 3 4 Osmoregulation Osmoregulation -Ionic and Osmotic Balance -life arose in salty sea -extracellular fluids ~ similar -Kidney Function -dist’n limited by temperature and osmotic pressure (dehydration, ionic composition) -terrestrial organisms (and their descendents) regulate internal environment (homeostasis) -salt and water regulation (waste excretion) -kidneys, salt glands, gills 5 6 1 Obligatory Osmotic Exchanges Obligatory Osmotic Exchanges 1-Gradients -Frog in freshwater -Fish in ocean 4-Feeding, Metabolism, Excretion -metabolic waste products ammonia, urea, etc. -metabolic water (desert!) -ingestion of salts -kidneys, salt glands, gills (more later) 2-Surface-to-Volume Ratio -Small animals dehydrate or hydrate more rapidly -Skin, and Respiratory surface (higher metabolism with higher per/gram respiratory surface) 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) 3-Integument Permeability -Transcellular or Paracellular -Aquaporins = water channel proteins -Frogs vs. Lizards, Pelvic Patch etc. 7 Osmoregulation Osmoregulation -Water Breathing -Water Breathing 1. Fresh Ambystoma tigrinum Blood osmolarity 200-300 mosm/L 2. Salt (~1,000 mosm/L) Most marine vertebrates hypo-osmotic (e.g., teleost or bony fishes) Water ~ 50 mosm/L - 8 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. - danger of losing water, gaining too many salts drink saltwater excess salts actively secreted (gills, kidneys) chloride cells for salt secretion 9 10 Osmoregulation -Air Breathing Have to lose water to allow gas exchange - Marine reptiles and marine birds can drink seawater and secrete salts in high [ ] Water Sources: - SALT GLANDS 1 Free - Mammals rely on kidney 2 Preformed 3 Metabolic (14-8) C6H12O6 + 6O2 ÅÆ 6CO2 + 6H20 11 12 2 Hill et al. 2004, Fig 26.15 Mouse-to-Elephant Curve (16-7) Allometry 4g shrew eats 2g/day elephant is 1 million x larger Allometry 13 14 Camel K-rat Lab rats K-rat Oryx Hopping Mouse 15 Osmoregulation ORGANS THAT CONTRIBUTE TO OSMOREGULATION IN VERTEBRATES 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 16 -Air Breathing Desert Mammals Behavior and Physiology (Eckert 14-9) 17 Kangaroo Rat -Reduce Activity -Remain in Cool Burrow -Humid -Water into dry seeds -Highly concentrated urine -Very dry feces (rectal absorption) -Metabolic water 18 3 Water Phyllomedusa Phyllomedusa sauvagi Lose water: evaporation urine feces salt glands eyes Eleutherodactylus coqui Pough et al., 2001 Alter behavior and physiology to minimize water loss Water balance limits activity in time and space Amphibs lose most water via evaporation - cutaneous resistance 1 dried mucus 2 cocoon 3 wax f t eo Rol abita h o r c mi 19 Pough et al., 2001 20 Water Chuckwalla Less evap. (lizards have more lipids in skin) osmoregulation mechanisms in gills, salt glands, and kidneys Monkey Tree Frog Anolis lizard Alligator Softshell Turtle More evap. Bufo, Spadefoots, Rana Pough et al., 2001 (free water surface) 21 22 Salt Secretion: Ionic & Osmotic Homeostasis recycle extracellular active intracellular Down electrochemical gradient (Paracellular) (Eckert 14-14) 23 24 4 Salt Glands Shark rectal glands to dispose of excess NaCl Figure in Box 26.2 Hill et al. 2004 Chloride Cell -blood hyperosmotic to seawater, but less salt -more urea and TMAO (trimethylamine oxide) -NaCl actively secreted 25 Shark Rectal Salt Glands 26 (Eckert 14-36) Salt Glands (Eckert 14-36) Salt-secreting cells: -Na/K-ATPase pump in basolateral membrane -generates gradient for Na+ by which Na+/2Cl-/K+ cotransporter drives up [Cl-] in cell -Cl- across apical membrane -Na+ follows paracellularly down electrochemical gradient (and H2O) -apical membrane impermeable to urea and TMAO -therefore iso-osmotic secretion with lots of NaCl Nasal/orbital salt glands of birds and reptiles -especially species in desert or marine environments. Hypertonic NaCl secretions (2-3x plasma osmolarity) Allows some birds to drink salt water and end up with osmotically free water … slightly different in birds and lizards Æ 27 28 Amblyrhynchus cristatus Fish Gills 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 (Eckert 14-14) 2 1 4 3 5 29 Hill et al. 2004, Fig 26.7 30 5 Freshwater fish: The mechanism basically reversed to allow uptake of salt from water against concentration gradient (recall lab paper on smolting) Sea ÅÆ Freshwater Switch between getting rid of excess salt in seawater and taking up salt in freshwater Na/K-ATPase to generate Na gradient proton pump to create electrical gradient Growth hormone and cortisol for Æ sea (more active chloride cells with more Na/K-ATPase activity) Prolactin for Æ freshwater (Eckert 14-31) 31 32 Gradients established and used…to move ions, water Osmoregulatory Mechanisms Apical surface (faces lumen and outside world) Basal surface (faces body and extracellular fluid) - Active movement of ions/salts requires ATP - Movement of water follows movement of ions/salts active passive (Eckert 14-12) (14-11) 33 Mammalian Kidney 34 6