© Cincinnati Zoo Homeostasis Maintaining a body in a “steady state” Environments where freshwater is scarce: • Desert • Invertebrates • Reptiles • Terrestrial Mammals • Ocean • Invertebrates • Fish • Marine Mammals, Turtles, & Birds • Hypersaline Lakes • Invertebrates World-wide distribution of desert environments •For example Shahara Desert is 3.5 mil sq. miles •Less than < 3 in. annual rainfall (6.7 cm) •Habitat: Rocky Plateaus (15%), Sandy Dunes (15%), Gravel Plains (70%) Adaptations for osmoregulation (water balance) • ↓ Water Loss – – – – Conservation Utilization (↑Efficiency) ↑ Storage capacity ↕ Solute/waste excretion LE 44-5 Water balance in a kangaroo rat (2 mL/day) Ingested in food (0.2 mL) Water balance in a human (2,500 mL/day) Ingested in liquid (1,500 mL) Ingested in food (750 mL) Water gain Derived from metabolism (1.8 mL) Feces (0.09 mL) Urine (0.45 mL) Derived from metabolism (250 mL) Feces (100 mL) Urine (1,500 mL) Water loss Evaporation (1.46 mL) Evaporation (900 mL) Adaptations for osmoregulation (water balance) • ↓ Water Loss – – – – Conservation Utilization (↑Efficiency) ↑ Storage capacity ↕ Solute/waste excretion LE 44-8 Proteins Nucleic acids Amino acids Nitrogenous bases —NH2 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 Waste management varies under different circumstances: • Habitat – Terrestrial vs. aquatic turtles • Reproductive strategy – Mammals: maternal transport of waste – Amphibians: diffuse Ammonia out of egg (lacking shell) – Birds/Reptiles: store as relatively less toxic uric acid • Diet – ↑ Animal tissue => ↑ N-Wastes • Metabolic requirement – ↑ metabolism => ↑ N-Wastes Adaptations for osmoregulation (water balance) • ↓ Water Loss – – – – Conservation Utilization (↑Efficiency) ↑ Storage capacity ↕ Solute/waste excretion • ↓Metabolic Requirements • Unique physiologic structures – Renal structure – Salt Glands Reduce Metabolic Requirements © Cincinnati Zoo – Reduce metabolic rate • Slow heart, blood flow • Reduces O2 consumption • Reduces body temp – Reduce activity 100 µm 100 µm Hydrated tardigrade Dehydrated tardigrade • Seek shelter, such as a burrow, shade, deeper water – e.g. Fennec, Leatherback • Enter torpor (or estivation, in summer) – resting state – e.g. Tarigrade, Mulgara, Longnecked Turtles Basic concept review: Diffusion Osmosis • Solutes move from greater solute concentration to lower conc. • Diffusion of water through a selectively permeable membrane Surface area to volume (SA:V) d=2 3:2 3:1 Chptr 40: Animal Form & Function, pp. 820-821 d=1 Kingdom Animalia Phylum Chordata Subphylum Vertebrata Mammalia Order Artiodactyla Family Camelidae Dromedary or Arabian Camel (Camelus dromedarius) Adaptations to ↓ Heat Loss • Anatomy – Hump (Fat = H20 Storage) – Lips (chewing tough vegetation) – Double eyelashes • Physiology – Increase body temp to match ambient temp – Body temp. range: 34 - 42°C (8°C range) • Behavior – Huddling – Tracking sun Class Water lost per day (L/100 kg body mass) LE 44-6 4 3 2 1 0 Control group (Unclipped fur) Experimental group (Clipped fur) Kingdom Animalia Phylum Chordata Subphylum Vertebrata Class Mammalia Order Carnivora Suborder Caniformia Family Canidae Fennec Fox (Fennecus zerda [=Volpes zerda]) Sahara Desert © Cincinnati Zoo • Anatomy – Small body size – Large ears – Furred foot pads • Behavior – Burrowing (↑ Humidity) – Nocturnal • Physiology (↓Water Loss) – Reduced heart rate – Reduced metabolic rate 85 - 95 F Banholzer, U. 1976. Water balance, metabolism, and heart rate in the Fennec. Naturwissenschaften 63 (4): 202-203. Kingdom Animalia Phylum Chordata Subphylum Vertebrata Class Mammalia Infraclass Marsupialia Order Dasyuromorphia Suborder Dasyuridae Family Dasyurinae Mulgara (Dasycercus cristicauda) • Occurs in arid, sandy regions of Australia • Related to marsupials • Example of convergent evolution with rodents – Small size, long tail – Fossorial (burrowing) – Nocturnal Bannertail kangaroo rat (Dipodomys spectabilis) –Torpor bouts 3 to 12 hr thus reduce metabolism to <12% of normal rate** *Nocon, W. 1999. "Dasycercus cristicauda" (On-line), Animal Diversity Web. Accessed May 24, 2008 at http://animaldiversity.ummz.umich.edu/site/accounts/information/Dasycercus_cristicauda.html. **Geiser, F and P. Masters. 1994. Torpor in relation to reproduction in the mulgara, Dasycercus cristicauda (Dasyuridae: Marsupialia) J. THERM. BIOL. 19 (1) pp. 33-40. LE 44-18c Roadrunner (Geococcyx californianus) Desert iguana (Dipsosaurus dorsalis) Marine Environment “Water, water everywhere, nor any drop to drink” - Coleridge • Ionic gradient set up: organism less salty than environment, salts want to enter body, water to leave (dehydration) • Most marine invertebrates are osmoconformers • Most marine vertebrates and some invertebrates are osmoregulators • Hypotonic – less salty than environment • Hypertonic - saltier than environment Marine Invertebrates Polyclad Flatworm • Most inverts are osmo – isotonic with environment (e.g. sponges) • Some have specialized protonephridia composed of ciliated flame cells to transport solutes and waste products for elimination Filtration Metanephridia: most annelids Malpighian tubules: insects Freshwater vs. Marine Fish Osmotic water gain through gills and body surface Gain of water and salt ions from food and seawater Osmotic water loss through gills and body surface water and some ions in food salt ions by gills Excrete large amounts of Dilute urine Freshwater animals are hyperosmotic to their environment They lose salts by diffusion and maintain water balance by excreting large amounts of dilute urine Salts lost by diffusion are replaced by foods and uptake across the gills Excretion of salt ions from gills Excretion of salt ions and small amounts of water in scanty urine from kidneys Marine bony fishes are hypoosmotic to sea water They lose water by osmosis and gain salt by diffusion and from food They balance water loss by drinking seawater Kingdom Animalia Phylum Chordata Subphylum Vertebrata Class Reptilia Order Testudines Family Cheloniidae Green Sea Turtle (Chelonia mydas) • The primary osmo-regulatory mechanism in sea turtles is the salt gland • The surface area: volume ratio is different for age classes: a 50g immature has greater surface area, and larger relative salt glands (0.3% of body size) than a 50 kg subadult (0.05 –0.1%) – i.e., osmotic challenge varies by age Kingdom Animalia Phylum Chordata Subphylum Vertebrata Class Reptilia Order Testudines Family Dermocheliidae • Salt glands secrete monovalent ions (Na+), which is the main constituent of seawater, while renal system processes bivalent ions (Mg++) Leatherback Sea • Sea turtles, marine reptiles & marine birds: super-saline secretions from salt glands Turtle (Dermochelys coriacea) – Lacrymal (turtles) – eye secretions – Nasal (lizards) – Post-orbital (birds) – Sublingual or premaxillary (snakes) – Lingual (crocodiles) Prange, H.D. 1985. Osmoregulation: water and salt balance in sea turtles Copeia 1985 (3): 771-776. Hudson, D. M. and P. L. Lutz 1986. Salt Gland Function in the Leatherback Sea Turtle, Dermochelys coriacea Copeia, 1986 (1):247-249 Salt glands in marine birds Nasal salt gland Nostril with salt secretions Pink-footed Shearwater (P. Hodum) Puffinus cretapus • Salt glands of marine birds remove excess sodium chloride from the blood • Use transport epithelia, which are specialized cells that regulate solute movement arranged in complex tubular networks Lumen of secretory tubule Vein Capillary Artery Secretory tubule NaCl Transport epithelium Direction of salt movement Blood flow Secretory cell of transport epithelium Central duct Structure of Kidney Kidney Renal medulla Renal cortex Ureter Urinary bladder Urethra Ureter Kidney structure Section of kidney from a rat Afferent arteriole Glomerulus from renal Bowman’s capsule artery Proximal tubule Peritubular capillaries Renal cortex SEM Collecting duct 20 µm Renal medulla Distal tubule Collecting duct Descending limb Nephron Loop of Henle Ascending limb Filtrate and blood flow LE 44-13 Vasa recta LE 44-14 Proximal tubule NaCl Nutrients HCO3– K+ H2O H+ NH3 Distal tubule H2O NaCl K+ HCO3– H+ CORTEX Descending limb of loop of Henle Filtrate H2O Salts (NaCl and others) HCO3– H+ Urea Glucose; amino acids Some drugs Thick segment of ascending limb NaCl H2O OUTER MEDULLA NaCl Thin segment of ascending limb Key Collecting duct Urea NaCl Active transport Passive transport INNER MEDULLA H2O Kidney: Nephron Kingdom Animalia Phylum Chordata Subphylum Vertebrata Class Mammalia Order Carnivora Suborder Pinnipedia Family Otariidae California Sea Lion (Zalophus californianus) • Derive water from food – fish, squid • All marine mammals have reniculate kidneys • which means that instead of having two single beanshaped kidneys, each kidney is instead made up of grapelike clusters of smaller, independent kidney units, or renicles, as shown Cross-section Marine Wildlife Veterinary Care & Research Center ADH: Anti-diuretic Hormone Hypersaline Environments Mono Lake, CA (HN) Often very simple food web & trophic level structure – Hypersaline lakes: – e.g. Mono Lake, Great Salt Lake American Avocet Great Salt Lake, Utah (photo: NASA) • Lower trophic level, low diversity: Brine shrimp (Artemia spp.) • Higher trophic level, great diversity: shorebirds, gulls & grebes http://ut.water.usgs.gov/shrimp/index.html "Brine Shrimp and Ecology of Great Salt Lake", United States Geological Survey Hypersaline Environments Avocets feeding in hypersaline Mono Lake, Lee County, CA 2007 © H. Nevins Hypersaline Environments Laysan Is., Hawaii (photo: USFWS) Laysan Ducks (photo: USFWS) – High level of endemism – Hypersaline lagoons: – e.g. Laysan Is., NW Hawaiian Islands • Brine flies (Ephydra spp.) • Laysan Duck (Anas laysanensis) READY FOR a QUIZ ??? California Gull, Mono Lake, CA 2007 © H. Nevins Quiz: • Name three taxa which occur in freshwater-scarce environments • Give a key physiological or behavioral adaptation for each to reduce water loss • How would you expect the structure of the loop of Henle to differ among a tropical forest and a desert rodent? Answers: • Name three species which occur in freshwaterscarce environments – Camelus, Artemia, Zalophus, others… • Give a key physiological or behavioral adaptation for each to reduce water loss – Camelus: hump - water storage, orient to sun, fur – Mulgara: fossorial, torpor – Zalophus: reticulate liver • How would you expect the structure of the Loop of Henle to differ among a tropical forest and a desert rodent? – Shorter in tropics, longer in desert to increase water absorption Questions? Avocets feeding in hypersaline Mono Lake, Lee County, CA 2007 © H. Nevins