osmoregulation by birds - ecology & evolutionary biologyeebweb.arizona.edu/courses/ecol437/ecol...
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Osmoregulation by Birds
Eldon J. BraunDepartment of PhysiologyUniversity of Arizona
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Maintenance of the Internal Environment
Osmoregulation means the maintenanceof the homeostasis of internal environment.
What constitutes the internal environment?
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Birds osmoregulated well
Birds inhabit all environments
Aquatic
Fresh water
Marine
Estuaries Terrestrial
Polar
Temperate
Desert
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In terms of osmoregulation, mammals are the unusual group
Kidneys are only osmoregulatory organ
Osmoregulation among other vertebrates
Fish, amphibians, reptiles, and birds
Multiple organs function in osmoregulation
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ORGANS THAT CONTRIBUTE TO OSMOREGULATION IN VERTEBRATES
Group Osmoregulatory Organs Fish Kidneys Gills Bladder Intestine Amphibians Kidneys Gills Bladder Skin Intestine Reptiles Kidneys Salt Glands Intestine Birds Kidneys Salt Glands Intestines Mammals Kidneys
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Osmoregulation by birds: Organs Involved
Kidneys
Lower gastrointestinal tract
Salt glands
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Avian salt glands
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The avian renal and gastrointestinal systems must function in concert in the regulation of ion and fluid balance.
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As birds do not have urinary bladders, the ureteral urine is refluxed from the cloaca into colon
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Evolutionary Rationale for this Type of Arrangement
Excess mass of urinary bladder
GFRs of Birds and Mammals Do NotDiffer
Fraction of Filtered Water Reabsorbed by Kidney
Less by Avian Kidney
Urine of Birds in Constant “Flux”
(i.e. urine entering lower GI tract)
Argument does not “hold water”
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Urine to plasma osmolar ratio
How well kidneys of animals concentrate urine isUsually expressed as the ratio of the urine osmolalityTo the plasma osmolality.
Or simply the U/Posm
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Birds or the avian kidney does not concentrate urine to a high degree
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Maximum U/Posms of some mammals
Values range from about 1 in the Mountain Beaver to ca. 25 in some of the small desert Rodents.
Humans U/Posm?
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Urine-to-Plasma Osmolar Ratios for Birds (U/Posm)
Ring-necked Pheasant 1.5Senegal Dove 1.7Savannah Sparrow 1.7King Quail 1.8White-crowned Sparrow 1.8Domestic Fowl 2.0Budgerigar 2.3House Finch 2.4Singing Honeyeater 2.4Stubble Quail 2.6
Mean 2.05
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Comparison of U/Posms between birds and mammals
Not valid comparison to make Urine in lower GI tract
End products of nitrogen metabolism Uric acid vs. urea
Effects of conc. fluid in lower GI tract
Urea ca. 50% of solutes in urineUric acid not in solution
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Plasma and urine osmolality of Desert Quail
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Nitrogen Excretion in Birds
Compound PercentUrea 4Ammonium 20Uric Acid 76
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Solubilites of Nitrogen-Containing CompoundsCompound Solubility (mmol/L)
Uric Acid 0.381Ammonium Urate 3.21
Sodium Urate 8.32Potassium Urate 14.75Urea 16,650
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Structure of Uric Acid
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Evolutionary Rationale for this Type of Arrangement(i.e. urine entering lower GI tract)
Crystal of Uric Acid
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Evolutionary Rationale for this Type of Arrangement(i.e. urine entering lower GI tract)
Physical form of uric acid in avian urine
Small spherical structures
Spheres ca. 65%uric acid
Uric acid boundTo a matrix protein
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Prevention of Sphere Coalescence
Protein in avian ureteral urine
Avian urine contains 5 mg/ml protein
Protein conc. in human urineca. 0.05 mg/ml
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Nature of Protein in Urine of BirdsSDS PAGE of avian Urine and plasma
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Energy in Avian Ureteral Urine
Male Female
Kcal/Day 5.3 12.4% BMR 5.4 11.3
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SDS PAGE of Avian Excreted Fluid
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Modification of Urine in Lower GI Tract of Birds
Anderson & Braun
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Degradation of Uric Acid in Lower GI Tract
68% of uric acid in ureteral urine
Bacterial action
• Fate of liberated nitrogeno Glutamic acid
Renal tubules--Buffer H ions Gluconeogensis Citric acid cycle
o Short chain volatile fatty acids
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Products Formed From the Breakdown of Uric Acid in Avian Lower GI tract
77% of [15N]uric acid introduced into ceca of cockerelsdisappeared in 60 min
And nitrogen appeared as ammonia and rapidly absorbed
Karasawa, 1989
Labelled nitrogen appeared in plasma within glutamine
Where do these product go?
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Vasculature Surrounding the Avian Kidney
Akester
Coccygomesenteric veindrains into renal portal system
Birds have a functional renal portal system
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Tubule Lumen Cells Peritubule Side
Blood fromrenal portalsystem
Glutamine
Glutamine
NH3NH3
Na+
H+H+
NH4+
Use of glutamine by renal tubules (To buffer hydrogen ions)
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Proteins
Amino Acids
Pyruvate
Acetyl CoA
Citric Acid cycle
Glyceraldehyde3-phosphate
Glucose
Fatty Acids
Ammonia
Possible Pathway for Utilization of Uric Acid
Also, deamination of glutamine produces ketoglutaric acid that can enter the krebs cycle
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Summary
Birds osmoregulate wellMultiple organ systems contribute to osmoregulation in birds
Salt glandsKidneysGastrointestinal tract
Uric acid as an end product to nitrogen metabolismForm of uric acid in urine“Value” of uric acidRecycling of the nitrogen of uric acid