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Osmoregulation
■ Ability to hold constant total electrolyte
and water content of the cells.
■ This is critical for both survival and
success.
■ Higher Vertebrates dependent on
regulation of internal electrolytes at 1%
or less
■ Tolerance for fluid volume even more
critical
Fluid Volume Comparisons
■ Hypoosmotic fluid with the lowest
concentration
■ Hyperosmotic fluid with the highest
concentration
■ Isoosmotic fluids of equal
concentration
Cellular Response to Variations in
Osmotic Pressure (tonicity)
■ Cells will respond to being placed in these
solutions
■ Hypotonic solution cells will take on fluid or
swell (possiblilty of lysis) when placed in this
fluid
■ Hypertonic cells will loose fluid or shrink
(crenation)
■ Isotonic cells will show not net change in
fluid volume
Cellular Response to Various Osmoticity Levels
Categories of Osmotic Exchange
■ Oligatory - animal has little if any control over
this category
■ Regulated - physiologically controlled and help
maintain homeostasis
Obligatory Exchanges
■ Body fluids are subdivided into two main
compartments intracellular and extracellular
■ Extracellular fluid is further subdivided into
interstitial fluid, blood plasma and transcellular
fluid
■ Transcellular fluid includes digestive secretions,
cerebrospinal fluid, synovial fluid, peritoneal
fluid, ocular fluid, and other fluids produced by
cells in special compartments
Human Example
Water Volume
■ Man and mammals in general are about 50
70% water by weight
■ Submammalian percentages are higher (70
80%) with more intracellular volumes
Intra and Extra Cellular Solutes
■ Predominant extracellular solutes include
NaCl, urea and TMAO
■ predominant intracellular solutes include
potassium, urea and TMAO
■ Osmolarity ranges from 2201000 mOsm with
most less than 350mOsm
■ Chondrichthyes and coelecanth maintain
osmotic pressures of about 1000 mOsm due
to retention of organics such as Urea
Variations is Solutes
Osmotic Problems Correlated With
Environment
■ sea water increased osmolarity due to
dissolved solvents which leads to water loss
and an accompanying high magnesium,
calcium and sulfate concentrations resulting
in ionic imbalance
■ fresh water dilution and loss of electrolytes
■ terrestrial dessicating environment, similar
to sea water but with no electrolyte
imbalances
Osmoregulatory Mechanisms
■ 1. Regulation of water entering the body
(alteation of skin and gill permeability, and
reduced or increased ingestion)
■ 2. Regulation of water loss (alteration of
surface permeability or excretory (renal) loss)
■ 3. Regulation of electrolyte loss (renal)
■ 4. Cutaneous uptake or excretion of ions
(usually oral or branchial)
■ 5. variation of osmotic pressure through
retention of organic solutes
Categories of Regulation
■ Osmoconformers many invertebrates and
only the hagfish as a vertebrate
– Although the hagfish does not truly regulate its
osmotic balance and conforms to that of the
ocean, it does regulate the concentration of ions
■ Osmoregulators
– Sodium and chloride are higher in blood than in
the ocean.
– Magnesium, calcium and sulfate are lower than in
the ocean.
– The kidney is responsible for this ionic regulation.
Elasmobranchs
■ Elasmobranchs are isoosmotic to their
environment due to the retention of organic
solutes (urea, TMAO) in their blood.
■ These are readily available due to biosyn
thetic pathways in the liver.
■ Excess salts are excreted by either the
kidney or the rectal gland.
Osmoregulatory Mechanisms - 1
Osmoregulatory Mechanisms - 2
Hypoosmotic Environment
■ Two major problems excess water and loss
of solutes
■ three solutions
– 1. decreased permeability or exterior surfaces
aquatic amphibian skin is 1/3 to 1/4 the
permeability of toad skin , semiaquatic skin is
intermediate
– 2. increased water excretion
– 3. increased salt uptake through epithelial
transport methods
Hypoosmotic Environment Continued
■ Contractile vacuoles in many microorganisms
eliminate excess water
■ Kidney excretion increased water loss due
to increased filtration and large volume of
urine production
■ In euryhaline fish these changes take place
as they migrate from salt water to FW and
vice versa
Contractile Vacuoles
When Paramecia are examined using the light microscope the contractile
vacuoles(CV) can be seen as a pair of spherical structures surrounded by a
series of radiating ducts. These ducts collect water and pump it into the central
area by means of sets of contractile fibers. In turn, the central sphere contracts
when it is full, forcing water to the outside of the protist.
Hyperosmotic Environment
■ three problems dessication, divalent ionic
imbalance and excess salt
– Teleosts drink sea water, absorb in gut (70
80%), desalinate with epithelial transport across
the gills
– Elasmobranchs retain urea and TMAO which is
also seen in crab eating frog of SE Asia
– Marine birds and reptiles eliminate salt through
nasal and or orbital glands and show reduced
urine flow to avoid dessication
– Mammals metabolic water and formation of a
concentrated urine (1 Kg of herring can yield up to
600 ml of water urine can be concentrated up to
5X)
Salt Glands in Sea Birds
Nasal Gland Function
Terrestrial
■ one main problem which is the loss of water
■ desert rodents produce urine at 1520X
plasma concentration normal rats 56X
■ Metabolic water production 1 g of fat can
yield 1.07 g of water, carbohydrate only 0.6g
and protein 0.4 g
■ the net yield after energy utilization is
greatest with carbohydrates
■ many animals also show behavioral
adaptations which reduce water loss
Desert Rodent Mechanism
Kidney- Primary Osmoregulatory Organ
■ Originally very simple a large glomerulus and
collecting duct used to regulate divalent ions
(hagfish)
■ FW fish large glomerulus for filtration, with
proximal and distal tubules specialized for
reabsorption of salt, glucose, and AA
■ Marine fish small glomeruli, with no distal
tubule which is used primarily to excrete
divalent ions
■ Elasmobranchs large glomeruli, complex
filtration and reabsorption of metabolites
Kidney Continued
■ Terrestrial used to reduce water loss and is
now the sole organ for for nitrogenous waste
removal
■ Reptiles have reduced glomeruli uric acid
secreted into the tubules (helps conserve
water)
■ Mammals large glomeruli for urea filtration
and a loop of henle added to assist in the
process of producing a highly concentrated
urine
Kidney Variations
In these groups the kidney is mostly for elimination of excess water.
Therefore the glomerulus is large and produces a high volume of filtrate.
Amphibian Kidney Function
■ The frog is also able to use its urinary bladder to aid water
conservation.
■ When in water, the frog's bladder quickly fills up with a
hypotonic urine.
■ On land, this water is reabsorbed into the blood helping to
replace water lost through evaporation through the skin.
■ The reabsorption is controlled by a hormone similar to
mammalian ADH.
Other Kidney Variations
Filter just enough to flush the uric
acid which is secreted into the
tubules to the cloacca where most
water is reabsorbed. Some reptiles
have no glomeruli at all. Most
Birds have a similar type of kidney
■ Determines the type of excretory product
produced
■ NH3 aquatic (ammonotelic) high toxicity,
needs much water to dilute
■ urea invertebrates and teleosts use de
gradation of uric acid to produce (uricolytic)
■ most vertebrates use the ornithine urea cycle
to produce urea (soluble)in the liver(ureotelic)
■ uric acid non toxic, insoluble insects, birds,
snakes, lizards (uricotelic)
Nitrogen Metabolism
Salt Glands of Birds
■ Found above eye opens into nasal cavity
■ Capable of producing fluid at 10002000
mOsm (SW=1000 mOsm/plasma 230 mOsm)
■ This allows them to generate FW by desalini
zation of sea water
■ The fluid output varies with the amount of
water ingested concentration constant
■ Increased efficiency over the kidney for ionic
excretion
■ Develops in response to increased salt active
transport system
Rectal Gland
■ salt enters by diffusion
■ concentrates to 1000 mOsm with 90% due to sodium
chloride
■ fluid production is sporadic
Gill Exretion
■ used for sodium chloride other ions handled
by the kidney
■ chloride cells in the gill filaments (secondary
lamellae) which are mitochondrial rich
(NaKATPase used)
■ chloride pumped against its concentration
gradient (2X sea water)
Summary of Fish (Euryhaline) Regulatory
Methods
■ The gill: Specialised 'chloride cells' secrete excess sodium
and chloride in a hyperosmotic environment such as
seawater.
■ The kidney: Renal tissue produces copious amounts of
dilute urine in a hyposmotic environment, such as
freshwater, and small amounts of isosmotic urine in a
hyperosmotic environment, such as seawater.
■ The gut: As the teleost adapts to an increase in
environmental salinity water is lost through the skin and
gills. To compensate the animal drinks the surrounding
medium.
Freshwater Adapted Fish Osmotic Balance
Chloride Cells
■ independent sodium and chloride transport
■ sodium is transported in the form of bicarbon
ate and chloride either as KCl or CaCl2
Kidney (Gross Structure)
■ Bean shaped hilus area from which ureter
leaves the renal sinus internally
■ Covered by the renal capsule
■ In cross section one can see the cortex and
medulla
■ Medulla is subdivided into 818 pyramids
which empty through renal papillae into the
renal sinus
Kidney
Nephron
■ Bowman’s Capsule structure which
encloses the glomerlus which is the network
of capillaries which carries out the active
filtration.
■ Blood enters the glomerulus by way of the
efferent arteriole
■ Smaller afferent arteriole exits the capsule
and forms the peritubular capillaries which
surrounds the rest of the nephron and in
cludes the vasa recta which is a counter
current system associated with the loop of
henle.
Glomerulus
Nephron Continued
■ Proximal Convoluted Tubules primar area of
reabsorption and secretion
■ Loop of Henle area of filtrate concentration
as it extends deep into the medullary area
■ Distal Convoluted Tubules area of further
reabsorption (primarily water)
■ Collecting Duct area of final water reab
sorption for determining the final
concentration of the urine
Nephron Structure
Nephron Processes
■ 1. Glomerular filtration due to high pressure in the
capillaries
– fluid in the Bowman’s
capsule has most sub
stances in concentra
tions equal to their
plasma concentrations
(proteins are an
exception)
– nonselective process
based strictly upon
molecular size and blood
pressure
Glomerular Filtration
– GFR = UinV/Pin (inulin used because it is
inert, 100% filtered and none is
reabsorbed)
– Clearance of inulin can be used as a
basis for calculating the clearance of other
substances.
– glucose clearance is less than 1% that of
inulin indicating reabsorption
– other substances may have a higher
clearance rate indicating secretion
Factors Affecting Filtration
■ 2. tubular reabsorption of water (99%) and
most salts, with most in the proximal tubule.
Some is active and some is passive due to
osmotic gradients.
■ 3. tubular secretion of certain excreted
compounds is a selective and active process
which is used mostly for potassium, hydrogen,
and bicarbonate as well as foreign substances
such as drugs
Nephron Processes
■ 4. tubular synthesis process such as
deamination of amino acids with NH3 then
diffusing into the lumen where it combines
with H+ to be excreted in the urine
Processes In Urine Formation
4. Synthesis
Tubular Reabsorption
■ most energy used in NaCl reabsorption
■ 2/3 of water in the proximal tubule
■ remaining reabsorbed by the distal tubule,
collecting ducts or loop of henle
■ waste products are poorly reabsorbed (urea
only 44%)
■ useful products nearly all resorbed (glucose)
some not regulated
Tubular Reabsorption
■ water and ions are usually regulated
reabsorption usually by either diffusion or
mediated transport
■ many reabsorbed substances have a
transport maximum glucose and Vitamin C
■ urea is passively reabsorbed the same is
true of most lipid soluble substances
■ the liver will sometimes convert substances
to polar metabolites which diffuse slowly and
thus increases their excretion
Sodium Reabsorption
Passive due to Na+ gradient
Sodium Reabsorption
Proton Exchange
Final Reabsorption of Water
■ Proximal tubules are highly permeable to
water
■ The reabsorption of water and sodium leads
to high chloride reabsorption as well
■ Permeability of the collecting ducts and distal
tubules can vary based upon the presence or
absence of ADH
■ ADH > activates CAMP > opens water
channels which leads to low urine volume
diabetes insipidus no ADH results in up to
■
25L of urine per day
Water Reabsorption
■ production of hyperosmotic urine 1400
mOsm/L 5X plasma of 300
■ urea, sulfate, phosphate and waste = 600
mOsm/day
■ 600 mOsm/day / 1400 mOsm/L = 0.444L/day
obligatory water loss
Loop of Henle
■ Loop of Henle function results in hyperosmotic
interstitial fluid in the medulla
■ Fluid enters at 300 mOsm ascending limb actively
pumps sodium and chloride into interstitial fluid
(cotransport)
■ Descending limb highly permeable to water so the
urine which travels up becomes hypoosmotic due to
osmotic movement of water out
■ this can later be adjusted through the action of ADH
leading to reabsorption in the distal tubule and collecting
duct based on the gradient established by the loop
Loop of Henle Function
Countercurrent
Multiplier
Countercurrent Multiplier
Sodium Reabsorption Regulation
■ Aldosterone if none present 15 g/day
■ Stimulates the production of Na+ channels
(also in sweat glands, LO, and salivary)
■ Regulated by the reninangiotensin system
■ Intrarenal baroreceptors and cells which
sense sodium and chloride concentration
■ In fish corticosterone regulates salt loss
through the gills
Renin-Angiotensin System
Potassium Regulation
■ K+ channels present in membrane of the late
distal tubule and collecting ducts
■ Most potassium diffuses out into lumen
■ Aldosterone can increase potassium reab
sorption through a simple reflex
■ Increased K+ > increased aldosterone>
increased secretion
■ Aldosterone secreting cells are sensitive to
potassium levels
pH regulation