Osmoregulatory Systems in Fishes Maintaining homeostasis with respect to solute concentrations and water content Homeostasis Chapter 7 Zoology 1450 Introduction Homeostasis = maintaining steady state equilibrium in the internal environment of an organisms Much is done involuntarily by action of hormones, enzymes and osmoregulatory processes. Although occasionally fish do just “pick up and move” if environmental conditions are unfavorable. Topics Osmoregulation Endocrine system Thermal regulation Definitions Homeostasis = maintaining steady state equilibrium in the internal environment of an organisms Solute homeostasis = maintaining equilibrium with respect to solute (ionic and neutral solutes) concentrations Water homeostasis = maintaining equilibrium with respect to the amount of water retained in the body fluids and tissues Definitions, continued Osmotic concentration - Total concentration of all solutes in an aqueous solution: measured in units of osmolals = 1 mole of solute/liter of water or milliosmolals = 1/1000th of one osmolal Osmoregulation in different environments Challenge to homeostasis depends on steady state concentration of solutes in the body fluids and tissues as well as concentration of solutes in the external environment marine systems: environment concentration = 34 36 parts per thousand salinity = 1000 mosm/l freshwater systems: environment concentration < 3 ppt = 1 - 10 mosm/l Osmoregulation in different environments Each species has a range of environmental osmotic conditions in which it can function: stenohaline - tolerate a narrow range of salinities in external environment - either marine or freshwater ranges euryhaline - tolerate a wide range of salinities in external environment - fresh to saline: short term changes: estuarine - 10 - 32 ppt, intertidal - 25 - 40 long term changes: diadromous fishes Four osmoregulatory strategies in fishes 1. Isosmotic (nearly isoionic, osmoconformers) 2. Isosmotic with regulation of specific ions 3. Hyperosmotic (fresh H20 fish) 4. Hyposmotic (salt H2O fish) Osmoregulation Strategies Osmoconforming (no strategy) Hagfish internal salt concentration = seawater. However, since they live IN the ocean....no regualtion required! Osmoregulation Strategies Elasmobranchs maintain internal salt concentration ~ 1/3 seawater, remaining 2/3 is urea and trimethylamine oxide (TMAO). So total internal osmotic concentration equal to seawater. Gill membrane has low permeability to urea so it is retained within the fish. Because internal inorganic and organic salt concentrations mimic that of their environment, passive water influx or efflux is minimized. Osmotic regulation by marine teleosts... ionic conc. approx 1/3 of seawater drink copiously to gain water Chloride cells eliminate Na+ and Clkidneys eliminate Mg++ and SO4= advantages and disadvantages? active tran. Saltwater teleosts: passive diff. H2O drink Na+, Cl- Na+, Cl- Mg++, SO4= Na+, Cl- Mg++, SO4= chloride cells kidneys active passive Chloride Cell fig 6.2: sea water PC pavement cell ClCl- Cl- + carrier Na+ Cl- K+ chloride cell internal PC Na+ Na+ Na+, Cl- gut accessory cell Na+ pump Na+ Na+ K+ ATPase mitochondria tubular system Osmotic regulation by FW teleosts Ionic conc. Approx 1/3 of seawater Don’t drink Chloride cells fewer, work in reverse Kidneys eliminate excess water; ion loss Ammonia & bicarbonate ion exchange mechanisms advantages and disadvantages? Freshwater teleosts: don’t drink active passive H2O Na+, Cl- Na+, Cl- Ion exchange pumps; beta chloride cells water kidneys Ion Exchange Mechanisms freshwater interior Na+ active ATP pump NH4+ or H+ Clactive ATP pump HCO3- gill membrane Freezing Resistance: What fishes might face freezing? hagfishes? isotonic marine elasmobranchs? isotonic freshwater teleosts? hypertonic marine teleosts? hypotonic Solution for Antarctic fish Macromolecular antifreeze compounds peptides (protein) glycopeptides (carbohydrate/protein) rich in alanine { molecules adsorb (attach) to ice crystal surface interfere with ice crystal growth (disrupt matrix) Why is this important??? ice ruptures cells; hinders osmoregulation What about rapid ion flux? Euryhaline Short-term fluctuations in osmotic state of environment, e.g. in intertidal zone or in estuaries where salinity can range from 10 to 34 ppt with the daily tidal cycle: these fish have both kinds of chloride cells when salinity is low, operate more like FW fishes when salinity is high, operate like marine fishes kidneys function only under low salinity conditions Euryhaline Diadromous fishes (spend part of life in salt water, part in freshwater – catadromous (migrate seaward) or anadromous (migrate up river) hormone-mediated changes associated with metamorphosis - convert from FW adaptations to SW or vice versa, depending on direction of migration What about stress?? Stressors (handling, sustained exercise such as escape from predator pursuit) cause release of adrenaline (epinephrine) - for mediating escape, etc. Adrenaline causes diffusivity of gill epithelium to increase (become “leaky” of water & ions) This accentuates the normal osmoregulatory challenge for FW or marine fishes How to reduce stress in stressed fishes? Minimize the osmotic challenge by placing fish in conditions that are isosmotic add salt to freshwater, e.g. in transporting fish or when exposing them to some other shortterm challenge dilute saltwater for same situation with marine species Thermoregulation in Fishes Temperature effects on fish Temperature exhibits the greatest influence on fish’s lives! Affects metabolism Affects digestion Signals reproductive maturation and behavior Fish are conformers (well, sort of...) Body temperature is that of the environment Each species has particular range of temperatures that they can tolerate and that are optimal Big difference between what you can tolerate and what you thrive in... Behavioral Thermoregulation in Fishes Although fish are ectotherms, they can alter their body temperature by moving to habitats with optimal temperature Hot Fishes Some fish can maintain body temperature greater than ambient - tunas, billfishes, relatives (nearly endothermic) Use retia (similar to rete mirable) in swimming muscles to conserve heat, exchange O2, etc. Red muscle is medial rather than distal Billfishes have warm brains - heat organ from muscles around eye Practical application You’re management decisions and actions must account for fish responses to temperature gradients and limitations Endocrine Systems of Fishes Pituitary Gland - Master Gland Linked with hypothalamus of brain Produces hormones that affect other endocrine tissues - indirect influence Produces hormones that affect nonendocrine tissues directly Pituitary Gland Indirect influence ACTH - adrenocorticotrophic hormone TH - thyrotrophic hormone stimulates interrenal tissue production of cortisol stimulate thyroid production of thyroxin (growth, metamorphosis-i.e. flounder) GTH- gonadotrophic hormone stimulates gonads to produce androgens/estrogens Pituitary Gland Effects non-endocrine tissues directly pigmentation - melanophore stimulating hormone (MSH) affects long-term control of color osmoregulation - prolactin, vasotocin controls fresh/saltwater systems growth – somatotrophic hormone stimulates > length, cell multiplication Thyroid Gland isolated follicles distributed in connective tissue along ventral aorta controls metabolic rate affects metamorphosis, maturation facilitates switch between fresh & salt water Gonads gamete and sex hormone production controls gametes maturation cause formation of secondary sex characteristics: color, shape, behavior in fish, several sex hormones also serve as pheromones - e.g. goldfish males respond to hormones released with ovulation Other endocrine tissues in fishes chromaffin tissues-located near kidneys & heart produce adrenaline/noradrenaline – “fight or flight” increases blood flow through gills, ventilation rate interrenal (inside kidney) tissues produce cortisol, cortisone - stress response hormones (reduce inflamation) Other endocrine tissues in fishes pancreatic islets produce insulin - controls glucose, glycogen metabolism (glucagon production) corpuscles of Stannius produce stanniocalcin - controls Ca+2 inflow at gills Immune System Introduction Obviously, the immune system is important in homeostatic processes. Immune systems of fish have two components: non-specific and specific. As we will see, both are involved in protecting fish from visible as well as invisible disease causing agents. Non-specific immunity Skin & Scales—specific solid layers of protection from pathological and chemical stressors. Mucus secretion—traps microorganisms; preventing entry into body cavity or circulation Macrophages (phagcytes) and cytotoxic cells— part of the inflamatory response which destroy pathogens within the body before they can do harm. Specific Immune Response More of an active response where an “invader” is detected and destroyed. Primary organs: kidney, thymus, spleen, intestine. Antigens—invading compounds which provoke an immune response. Source: Cancer Research Institute (2002) www.cancerresearch.org/immhow.html Specific immune response: What if something does get in?? White blood cells called B lymphocyte cells (B cells) and T lymphocyte cells (T cells)—bind to foreign cells and begin replication and attachement to (sort of markers for things to come...). Occasionally, invader actually goes trough a macrophage first...then B cell responds Once B cells replicate, antibodies are produced which bind specifically to pathogens and tag them for destruction (eating) by macrophages! “Looks like meat’s back on the menu boys!!!” Questions???