1. Osmoregulatory Systems in Fishes Maintaining homeostasis with respect to solute concentrations and water content

2. Homeostasis Chapter 7 Zoology 1450

3. Introduction Homeostasis = maintaining steady state equilibrium in the internal environment of an organisms 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. 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.

4. Topics Osmoregulation Osmoregulation Endocrine system Endocrine system Thermal regulation Thermal regulation

5. Definitions Homeostasis = maintaining steady state equilibrium in the internal environment of an organisms 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 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 Water homeostasis = maintaining equilibrium with respect to the amount of water retained in the body fluids and tissues

6. 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 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

7. Osmoregulation in different environments Challenge to homeostasis depends on Challenge to homeostasis depends on steady state concentration of solutes in the body fluids and tissues as well as steady state concentration of solutes in the body fluids and tissues as well as concentration of solutes in the external environment concentration of solutes in the external environment marine systems: environment concentration = 34 - 36 parts per thousand salinity = 1000 mosm/l marine systems: environment concentration = 34 - 36 parts per thousand salinity = 1000 mosm/l freshwater systems: environment concentration < 3 ppt = 1 - 10 mosm/l freshwater systems: environment concentration < 3 ppt = 1 - 10 mosm/l

8. Osmoregulation in different environments Each species has a range of environmental osmotic conditions in which it can function: 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 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: euryhaline - tolerate a wide range of salinities in external environment - fresh to saline: short term changes: estuarine - 10 - 32 ppt, intertidal - 25 - 40 short term changes: estuarine - 10 - 32 ppt, intertidal - 25 - 40 long term changes: diadromous fishes long term changes: diadromous fishes

9. Four osmoregulatory strategies in fishes 1. Isosmotic (nearly isoionic, osmoconformers) 2. Isosmotic with regulation of specific ions 3. Hyperosmotic (fresh H 2 0 fish) 4. Hyposmotic (salt H 2 O fish)

10. Osmoregulation Strategies Osmoconforming (no strategy) Hagfish internal salt concentration = seawater. However, since they live IN the ocean....no regualtion required!

11. 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. 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. 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.

12. ionic conc. approx 1/3 of seawater ionic conc. approx 1/3 of seawater drink copiously to gain water drink copiously to gain water Chloride cells eliminate Na + and Cl - Chloride cells eliminate Na + and Cl - kidneys eliminate Mg ++ and SO 4 = kidneys eliminate Mg ++ and SO 4 = advantages and disadvantages? Osmotic regulation by marine teleosts...

13. Saltwater teleosts: drink active tran. passive diff. Na +, Cl - Mg ++, SO 4 = H2OH2OH2OH2O Na +, Cl - chloride cells Mg ++, SO 4 = kidneys

14. Chloride Cell fig 6.2 : pavementcell PC PC active passive chloride cell accessorycell sea water internal mitochondria tubular system Na + K+K+K+K+ Na + K + ATPase Na +, Cl - Na + Cl - gut carrier Na + pump +

15. Ionic conc. Approx 1/3 of seawater Ionic conc. Approx 1/3 of seawater Don’t drink Don’t drink Chloride cells fewer, work in reverse Chloride cells fewer, work in reverse Kidneys eliminate excess water; ion loss Kidneys eliminate excess water; ion loss Ammonia & bicarbonate ion exchange mechanisms Ammonia & bicarbonate ion exchange mechanisms advantages and disadvantages? Osmotic regulation by FW teleosts

16. Freshwater teleosts: active passive H2OH2OH2OH2O Na +, Cl - don’tdrink water kidneys Ion exchange pumps; beta chloride cells

17. Ion Exchange Mechanisms gill membrane freshwater interior activepump activepump Na + Cl - NH 4 + or H + HCO 3 - ATP ATP

18. Freezing Resistance: What fishes might face freezing? What fishes might face freezing?hagfishes?isotonic marine elasmobranchs? isotonic freshwater teleosts? hypertonic marine teleosts? hypotonic

19. Solution for Antarctic fish Macromolecular antifreeze compounds Macromolecular antifreeze compounds peptides (protein) peptides (protein) glycopeptides glycopeptides (carbohydrate/protein) (carbohydrate/protein) { rich in alanine molecules adsorb (attach) to ice crystal molecules adsorb (attach) to ice crystal surface surface interfere with ice crystal growth interfere with ice crystal growth (disrupt matrix) (disrupt matrix) Why is this important??? Why is this important??? ice ruptures cells; hinders osmoregulation ice ruptures cells; hinders osmoregulation

20. 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: 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 these fish have both kinds of chloride cells when salinity is low, operate more like FW fishes when salinity is low, operate more like FW fishes when salinity is high, operate like marine fishes when salinity is high, operate like marine fishes kidneys function only under low salinity conditions kidneys function only under low salinity conditions

21. Euryhaline Diadromous fishes (spend part of life in salt water, part in freshwater – catadromous (migrate seaward) or anadromous (migrate up river) 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 hormone-mediated changes associated with metamorphosis - convert from FW adaptations to SW or vice versa, depending on direction of migration

22. What about stress?? Stressors (handling, sustained exercise such as escape from predator pursuit) cause release of adrenaline (epinephrine) - for mediating escape, etc. 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) Adrenaline causes diffusivity of gill epithelium to increase (become “leaky” of water & ions) This accentuates the normal osmoregulatory challenge for FW or marine fishes This accentuates the normal osmoregulatory challenge for FW or marine fishes

23. How to reduce stress in stressed fishes? Minimize the osmotic challenge by placing fish in conditions that are isosmotic 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 short- term challenge add salt to freshwater, e.g. in transporting fish or when exposing them to some other short- term challenge dilute saltwater for same situation with marine species dilute saltwater for same situation with marine species

24. Thermoregulation in Fishes

25. Temperature effects on fish Temperature exhibits the greatest influence on fish’s lives! Temperature exhibits the greatest influence on fish’s lives! Affects metabolism Affects metabolism Affects digestion Affects digestion Signals reproductive maturation and behavior Signals reproductive maturation and behavior

26. Fish are conformers (well, sort of...) Body temperature is that of the environment Body temperature is that of the environment Each species has particular range of temperatures that they can tolerate and that are optimal 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... Big difference between what you can tolerate and what you thrive in...

27. Behavioral Thermoregulation in Fishes Although fish are ectotherms, they can alter their body temperature by moving to habitats with optimal temperature Although fish are ectotherms, they can alter their body temperature by moving to habitats with optimal temperature

28. Hot Fishes Some fish can maintain body temperature greater than ambient - tunas, billfishes, relatives (nearly endothermic) 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 O 2, etc. Use retia (similar to rete mirable) in swimming muscles to conserve heat, exchange O 2, etc. Red muscle is medial rather than distal Red muscle is medial rather than distal Billfishes have warm brains - heat organ from muscles around eye Billfishes have warm brains - heat organ from muscles around eye

29. Practical application You’re management decisions and actions must account for fish responses to temperature gradients and limitations You’re management decisions and actions must account for fish responses to temperature gradients and limitations

30. Endocrine Systems of Fishes

31. Pituitary Gland - Master Gland Linked with hypothalamus of brain Linked with hypothalamus of brain Produces hormones that affect other endocrine tissues - indirect influence Produces hormones that affect other endocrine tissues - indirect influence Produces hormones that affect non- endocrine tissues directly Produces hormones that affect non- endocrine tissues directly

32. Pituitary Gland Indirect influence Indirect influence ACTH - adrenocorticotrophic hormone ACTH - adrenocorticotrophic hormone stimulates interrenal tissue production of cortisol stimulates interrenal tissue production of cortisol TH - thyrotrophic hormone TH - thyrotrophic hormone stimulate thyroid production of thyroxin (growth, metamorphosis-i.e. flounder) stimulate thyroid production of thyroxin (growth, metamorphosis-i.e. flounder) GTH- gonadotrophic hormone GTH- gonadotrophic hormone stimulates gonads to produce androgens/estrogens stimulates gonads to produce androgens/estrogens

33. Pituitary Gland Effects non-endocrine tissues directly Effects non-endocrine tissues directly pigmentation - melanophore stimulating hormone (MSH) pigmentation - melanophore stimulating hormone (MSH) affects long-term control of color affects long-term control of color osmoregulation - prolactin, vasotocin osmoregulation - prolactin, vasotocin controls fresh/saltwater systems controls fresh/saltwater systems growth – somatotrophic hormone growth – somatotrophic hormone stimulates > length, cell multiplication stimulates > length, cell multiplication

34. Thyroid Gland isolated follicles distributed in connective tissue along ventral aorta isolated follicles distributed in connective tissue along ventral aorta controls metabolic rate controls metabolic rate affects metamorphosis, maturation affects metamorphosis, maturation facilitates switch between fresh & salt water facilitates switch between fresh & salt water

35. Gonads gamete and sex hormone production gamete and sex hormone production controls gametes maturation controls gametes maturation cause formation of secondary sex characteristics: color, shape, behavior 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 in fish, several sex hormones also serve as pheromones - e.g. goldfish males respond to hormones released with ovulation

36. Other endocrine tissues in fishes chromaffin tissues-located near kidneys & heart chromaffin tissues-located near kidneys & heart produce adrenaline/noradrenaline – “fight or flight” produce adrenaline/noradrenaline – “fight or flight” increases blood flow through gills, ventilation rate increases blood flow through gills, ventilation rate interrenal (inside kidney) tissues interrenal (inside kidney) tissues produce cortisol, cortisone - stress response hormones (reduce inflamation) produce cortisol, cortisone - stress response hormones (reduce inflamation)

37. Other endocrine tissues in fishes pancreatic islets pancreatic islets produce insulin - controls glucose, glycogen metabolism (glucagon production) produce insulin - controls glucose, glycogen metabolism (glucagon production) corpuscles of Stannius corpuscles of Stannius produce stanniocalcin - controls Ca +2 inflow at gills produce stanniocalcin - controls Ca +2 inflow at gills

38. Immune System

39. Introduction Obviously, the immune system is important in homeostatic processes. Obviously, the immune system is important in homeostatic processes. Immune systems of fish have two components: non-specific and specific. 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. As we will see, both are involved in protecting fish from visible as well as invisible disease causing agents.

40. Non-specific immunity Skin & Scales—specific solid layers of protection from pathological and chemical stressors. Skin & Scales—specific solid layers of protection from pathological and chemical stressors. Mucus secretion—traps microorganisms; preventing entry into body cavity or circulation 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. Macrophages (phagcytes) and cytotoxic cells— part of the inflamatory response which destroy pathogens within the body before they can do harm.

41. Specific Immune Response More of an active response where an “invader” is detected and destroyed. More of an active response where an “invader” is detected and destroyed. Primary organs: kidney, thymus, spleen, intestine. Primary organs: kidney, thymus, spleen, intestine. Antigens—invading compounds which provoke an immune response. Antigens—invading compounds which provoke an immune response. Source: Cancer Research Institute (2002) www.cancerresearch.org/immhow.html

42. 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...). 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 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! Once B cells replicate, antibodies are produced which bind specifically to pathogens and tag them for destruction (eating) by macrophages!

44. “Looks like meat’s back on the menu boys!!!”

45. Questions???