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Osmoregulatory and Excretory Systems

■ Ability to hold constant total electrolyte 
and water content of the cells.   
■ This is critical for both survival and 
■ Higher Vertebrates dependent on 
regulation of internal electrolytes at 1% 
or less
■ Tolerance for fluid volume even more 
Fluid Volume Comparisons

■ Hypoosmotic ­ fluid with the lowest 
■ Hyperosmotic ­ fluid with the highest 
■ Isoosmotic ­ fluids of equal 
Cellular Response to Variations in
Osmotic Pressure (tonicity)
■ Cells will respond to being placed in these 
■ Hypotonic solution ­ cells will take on fluid or 
swell (possiblilty of lysis) when placed in this 
■ Hypertonic ­ cells will loose fluid or shrink 
■ 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

■ Gradients between animal and environment - depends

upon animals surface are, degree of gradient and
permeability of animals surface
■ Small animals tend to dehydrate more rapidly than larger
animals due to reduced surface area:volume ratios
■ Nature of the integument can alter water loss or gain as
■ Feeding, metabolism and excretory mechanisms alter
water balance
■ Water is often lost due to respiratory mechanisms
Insect Cuticle
Respiratory Balance
Body Fluid Compartments

■ Body fluids are subdivided into two main 
compartments ­ intracellular and extracellular
■ Extracellular fluid is further subdivided into 
interstitial fluid, blood plasma and transcellular 
■ 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
■ 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 
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 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 
–  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 
Salt Glands in Sea Birds
Nasal Gland Function

■ 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 
■ 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 
■  Reptiles ­ have reduced glomeruli ­ uric acid 
secreted into the tubules (helps conserve 
■ Mammals ­ large glomeruli for urea filtration 
and a loop of henle added to assist in the 
process of producing a highly concentrated 
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 amphibian kidney also functions chiefly as a device

for excreting excess water. The permeable skin of the frog
provides an easy route for the fresh water of its pond to
enter by osmosis. But, as their name suggests, amphibians
also spend time on land. Then the problem is to conserve
water, not eliminate it.

■ The frog adjusts to the varying water content of its

surroundings by adjusting the rate of filtration at the
glomerulus. When blood flow through the glomerulus is
restricted, a renal portal system is present to carry away
materials reabsorbed through the tubules.
Amphibian Bladder

■ The frog is also able to use its urinary bladder to aid water
■ 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

Living in constant danger of

dehydration by the hypertonic
sea, there is no reason to pump
out large amounts of nephric
filtrate at the glomerulus. The
less water placed in the tubules,
the less that has to be
reabsorbed. So it is not
surprising that many bony fishes
have small glomeruli and some
have no glomeruli at all
Nitrogen Metabolism

■ Determines the type of excretory product 
■ 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 
■ Develops in response to increased salt active 
transport system
Rectal Gland
■ salt enters by diffusion
■ concentrates to 1000 mOsm with  90% due to sodium 
■ 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
■ The gill: Specialised 'chloride cells' secrete excess sodium
and chloride in a hyperosmotic environment such as
■ 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
Freshwater Adapted Fish Osmotic Balance
Chloride Cells

In freshwater adapted teleosts the chloride cells remove excess ions

such as bicarbonate and ammonium, and the Na, K-ATPase pump on
the basolateral membrance works to create an ionic gradient favoring
the movement of Na+ and Cl- into the cell from the external
Marine Adapted Fish Osmotic Balance
Chloride Cells

In seawater adapted euryhaline teleosts the chloride cells

are larger, and their principle role is to remove excess
Na+ and Cl- ions. Consequently, the Na, K-ATPase
pump works to create a concentration gradient favouring
the movement of these ions into the external
Amphibian Skin and Bladder

■ 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 is subdivided into 8­18 pyramids 
which empty through renal papillae into the 
renal sinus
■ Bowman’s Capsule ­ structure which 
encloses the glomerlus which is the network 
of capillaries which carries out the active 
■ 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 
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 
– fluid in the Bowman’s                                             
capsule has most sub                                                   
­stances in concentra­                                                   
     tions equal to their                                                    
         plasma concentrations                                         
            (proteins are an                                                 
– non­selective process                                             
based strictly upon                                               
molecular size and blood                                           
Glomerular Filtration

– GFR = UinV/Pin  (inulin used because it is 
inert, 100% filtered and none is 
–   Clearance of inulin can be used as a 
basis for calculating the clearance of other 
– glucose clearance is less than 1% that of 
inulin indicating reabsorption
– other substances may have a higher 
clearance rate indicating secretion
Factors Affecting Filtration

■ Net hydrostatic pressere difference between

the lumen of the glomerular capillaries and
the lumen of Bowman’s capsule
■ The colloid osmotic pressure of the blood
plasma (opposes filtration)
■ The hydraulic permeability of the tissue
separating the two compartments
Net Hydrostatic Pressure
Nephron Processes

■ 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 
■ 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 
■ 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 
■  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
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

■ Acid secreting (A) and base secreting (B)

cells in the distal tubule can help regulate
body pH
■ Buffering of renal filtrate by H2PO4- and
NH4+ permits greater secretion of protons
A-type cells
B-type Cells
Renal Filtrate Buffering
Control of Water Reabsorption

■ Final urine concentration determined

through alteration of the permeability of the
collecting duct
■ This is achieved through action of the
hormone ADH
■ Decreased blood pressure will stimulate
ADH production which will increase
permeability of the collecting ducts to water
ADH Regulation of Water Retention