OSMOREGULATION AND EXCRETION

● Homeostasis requires osmoregulation

○ Osmoregulation - the general term for the processes by which animals control solute
concentrations and balance water gain and loss
● Freshwater animals face a distinct challenge: watery environment threatens to dilute their body fluids
○ Conserve solutes and absorb salts to survive
● Animals must deal with ammonia in safeguarding their internal fluids
○ Ammonia - toxic metabolite produced by the dismantling of nitrogenous molecules (proteins and
nucleic acids)
● Excretion - process of ridding the body of nitrogenous metabolites and other metabolic waste products

Osmoregulation Balances the Uptake and Loss of water and Solutes

● Regulating the chemical composition of body fluids depends on balancing the uptake and loss of water
and solutes
○ Excessive uptake of water = swell and bust (lysis)
○ Excessive water loss - shrivel and die (shrivel)
● Driving force for the movement of water and solutes is a concentration gradient of one or more
solutes across a plasma membrane

Osmosis and Osmolarity

● Water enters and leaves cells by osmosis
○ Occurs when two solutions separated by a membrane
differ in total solute concentration
● Osmolarity - unit of measurement for solute concentration
○ No. of moles of solute per liter of solution
○ Human blood - 300 milliosmoles per liter (mOsm/L)
○ Seawater - 1,000 mOsm/L

Isoosmotic Hyperosmotic Hypoosmotic

● Solutions with the same ● The solution with the ● The solution that is more
osmolarity higher concentration of dilute (more water)
solutes

● Water molecules continually ● Water flows by osmosis ● Water flows by osmosis

cross the membrane at to here from here to
equal rates in both hyperosmotic sol’n
directions (with selectively (reducing concentration
permeable membrane gradient)
separating sol’ns)
● No net movement of water
by osmosis

Osmoregulatory Challenges and Mechanisms

● Two ways to maintain water balance:
1.) Osmoconformer
● To be isoosmotic with its surroundings
● All are marine animals
○ Living in waters that are stable in composition (constant internal osmolarity)
● No tendency to gain or lose water
2.) Osmoregulator
 ● To control internal osmolarity independent of that of the external environment
● Enables animals to live in environments that are uninhabitable for osmoconformers
○ Freshwater and terrestrial habitats, or move between marine and freshwater
environments
● Hypoosmotic environment - osmoregulator discharges excess water
● Hyperosmotic environment - osmoregulator takes in water to offset osmotic loss
● Stenohaline animals
○ An animal that cannot tolerate substantial changes in external osmolarity
● Euryhaline animals
○ Can survive large fluctuations in external osmolarity
○ Euryhaline osmoconformers - e.g. barnacles and mussels (alternately exposed to fresh and salt
water)
○ Euryhaline osmoregulators - e.g. striped bass and various species of salmon

Marine Animals Freshwater Animals

● Most marine animal invertebrates are ● Osmoregulatory problems of freshwater

osmoconformers animals are opposite of those of marine
○ Osmolarity is the same as that of animals
seawater ● Body fluids of freshwater animals must be
○ No substantial challenges in water hyperosmotic because animal cells cannot
balance tolerate salt concentrations as low as that
● Actively transport specific solutes that of lake/river water
establish levels in hemolymph different ● Face the problem of gaining water by osmosis
from those in the ocean b/c they have internal fluids w/ higher
○ E.g. Atlantic lobster (Homarus osmolarity than surroundings
americanus) maintain magnesium ion
concentration of less than 9mM vs. Solution:
50mM in their env ● Water balance relies on excreting large
amounts of very dilute urine and drinking
Osmoregulatory challenges (marin vertebrates) almost no water
● Address challenges of strongly dehydrating ● Salts lost by diffusion and in urine are
env. replenished by eating and by salt uptake
● Hypoosmotic to env. (generally) across their gills
1.) Marine “bony fishes” (+ ray-finned and lobe-
finned fishes, cod) Euryhaline fishes
● Internal salt concentration lower than ● Salmon and other euryhaline fishes that
seawater migrate between freshwater and seawater
● Constantly lose water by osmosis undergo dramatic changes in osmoregulatory
● Balance water loss by drinking a lot of status
seawater ● When in Rivers/streams (freshwater)
● Excess salts ingested with seawater ○ Osmoregulate like other freshwater
are eliminated through gills and fishes
kidneys ○ Produce large amts of dilute urine and
take up salt from dilute env. Through
● Osmoregulation is frequently coupled to gills
elimination of nitrogenous waste ● When move to the ocean
products such as urea ○ Acclimatize; become like bony fishes in
○ Elimination is important b/c high salt water
concentrations of urea can ○ Produce more of steroid hormone
denature (unfold) proteins and cortisol which increases the no. and
disrupt cellular functions size of specialized salt-secreting cells
2.) Sharks and other Chondrichthyans ○ Salmon in salt water then excrete
● Have an internal salt concentration excess salt from gills and produce
 lower than that of seawater (like bony only small amts of urine
fishes)
● Salt tends to diffuse from water into
their bodies (esp. Across their gills)
● Have trimethylamine oxide (TMAO)
produced by shark tissues
○ Combines with salts, urea, and
other compounds which result in
solute concentration in shark
tissues higher than
1,000mOsm/L
○ Thus, water slowly enters
shark’s body by osmosis and
in food (they don’t drink)

● TMAO also function as protection of

proteins from denaturing effect of
urea hence they have a high
concentration of urea in their body
(vs. bony fishes) → become
hyperosmotic to seawater
● Small influx of water into shark’s body is
disposed of in urine produced by
kidneys
● Urine - removes some salt that
diffuses into shark’s body; rest is lost in
feces or secreted from specialized
gland

Animals Living in Temporary Waters

● Extreme dehydration or desiccation is fatal for most animals
● Solution:
○ Few aquatic invertebrates that live in temporary ponds and in films of water around soil particles
can lose almost all their body and survive
○ Anhydrobiosis - Adaptation where animal enters dormant state when their habitats dry up
■ “Life without water”
■ Found in tardigrades or water bears - tiny invertebrates less than 1mm long
● Contain abt 85% water by weight in active, hydrated state
 ● Contain and dehydrate to less than 2% water and survive in inactive state, dry as
dust, or a decade or more
○ Just add water and within hours they’ll be rehydrated and moving
● Anhydrobiosis reqs. Adaptations that keep cell membranes intact
○ Contain large amounts of sugars
■ Disaccharide called trehalose protect the cells by replacing water that is normally
associated with proteins and membrane lipids
● Also used by insects that survive freezing in winter as well as plants resistant to
dessication

Land Animals
● Threat of dehydration is a major regulatory problem for
terrestrial plants and animals
● Adaptations that reduce water loss are key to
survival on land
○ Body coverings of most terrestrial animals
help prevent dehydration (lik waxy cuticle of
plants)
■ Insect exoskeleton
■ Shells of land snails
■ Layers of dead keratinized skin cells of
most terrestrial vertebrates (+ humans)
■ Nocturnal desert-dwellers to reduce
evaporative water loss b/c of lower temp
and higher humidity of night air
● Despite these adaptations, most terrestrial animals lose
water through many routes:
○ Urine
○ Feces
○ Across skin
○ Surfaces of gas exchange organs
● Land animals maintain water balance by drinking and
eating moist foods and by producing water
metabolically through cellular respiration

Energetics of Osmoregulation
● Maintaining an osmolarity difference between an animal’s body and its external env. Carries an energy
cost
● Diffusion tends to equalize concentrations in a system so osmoregulators must expend energy to
maintain the osmotic grandients that cause water to move in or out
○ Do so by active transport to manipulate solute concentrations in their body fluids
● Energy cost of osmoregulation depends on:
○ How different an animal’s osmolarity is from its surroundings (the higher, the more energy)
○ How easily water and solutes can move across animal’s surface
○ How much work is required to pump solutes across the membrane
● Energy cost to an animal maintaining water and salt balance is minimized by having body fluids adapted
to salinity of animal’s habitat
 ○ Body fluids of most animals in fresh water have lower solute concentrations than the body fluids
of their closest relatives in seawater
■ E.g. marine molluscs - body fluid concentration at 1,000mOsm/L; freshwater molluscs - 40
mOsm/L
○ Minimizing osmotic difference between body fluids and the surrounding env. Decreases the
energy cost of osmoregulation

Transport Epithelia in Osmoregulation

● Ultimate function of osmoregulation is to control solute
concentration in cells
○ Done indirectly by managing solute concentration of an
internal body fluid that bathes the cells
○ Insects and open circ. system - hemolymph (fluid
surrounding cells; like blood)
Vertebrates and closed syst. - interstitial fluid that
contains a mixture of solutes controlled indirectly by blood
○ Maintenance of composition of fluids depend on
structures ranging from individual cells that regulate solute
movement to complex organs such as the vertebrate
kidney
● Most animals, osmoregulation and metabolic waste disposal
rely on transport epithelia
○ Transport epithelia
■ One or more layers of epithelial cells specialized
for moving particular solutes in controlled amts in
specific directions
■ Typically arranged into tubular networks with
extensive surface areas
■ Face outside env. Directly or line channels connected to the outside by an opening on body
surface
■ Enables the albatross and other marine birds to survive on seawater
● Fluid drips from the tip of their beaks (concentrated sol’n of NaCl)
● Nasal salt glands
○ Packed with transport epithelia
○ Also found in sea turtles and marine iguanas
○ Use active transport of ions to secrete fluid much saltier than the ocean
○ Enable marine vertebrates to achieve a net gain of water
■ Also function in disposal of metabolic wastes

Nitrogenous Wastes
● Most metabolic wastes must be dissolved in water to be excreted from the body
○ Type and quantity of waste products may have a large impact on animal’s water balance
○ Waste products include: Nitrogenous breakdown products of proteins and nucleic acids
■ When proteins and nucleic acids are broken apart for energy or converted to carbs or fats,
enzymes remove nitrogen in the form of ammonia (NH3)
■ Ammonia - very toxic b/c its ion, ammonium, (NH4+) can interfere with oxidative
phosphorylation

Forms of Nitrogenous Waste

Ammonia Urea Uric Acid

Most aquatic animals (+ bony Mammals, most amphibians, Birds & many reptiles, insects, land
fishes) sharks, some bony fishes (marine) snails (animals that lay eggs)

● Animals that excrete ● Excreted by those who don’t ● White precipitate, usually
ammonia need access to a have access to sufficient mixed with feces (e.g. bird
lot of water water guano)
● B/c ammonia can be ● Main advantage: ● Relatively nontoxic
tolerated only at very low ○ Low toxicity ● Does not readily dissolve in
concentrations ● Main disadvantage: H2O
● Highly soluble ammonia ○ Reqs a lot of ● Excreted as semisolid
molecules (interconverting energy to be paste with very little H2O
between NH3 and NH4+) produced loss
easily pass through In vertebrates: ● Requires a lot of ATP for
membranes and are ● Urea is the product of an synthesis from ammonia
readily lost by diffusion to energy-consuming ● Humans and other animals
the surrounding water metabolic cycle that (not primarily uric acid
● In many invertebrates, combines ammonia with producers) generate a small
ammonia release occurs CO2 in liver amt of uric acid from
across the whole body In amphibians: metabolism
surface ● Switch between excreting ○ Diseases altering
ammonia in water (save this process causes
energy) and excreting urea uric acid crystals (in
(reduce excretory H2O loss) bladder for dogs or
gout in adult
humans, esp. males)

Most toxic Less toxic than Ammonia Least toxic

● Can only be safely
transported through and
excreted from body in large
volumes of very dilute sol’ns

Requires the most amount of water Requires less water Requires least amt of H2O

Requires least energy Requires more energy Requires the most energy

Influence of Evolution and Environment on Nitrogenous Wastes

● Type and amt of nitrogenous waste a species produces are matched to its env.
● Influence of availability of H2O:
○ Terrestrial turtles (dry areas) - uric acid
○ Aquatic turtles - urea and ammonia
● In some cases: Animal’s egg is the immediate env. Of relevance to the type of nitrogenous waste
excreted
○ Amphibian egg (no shell) - ammonia or urea can simply diffuse out of the egg
○ Mammal - soluble wastes produced by embryo can be carried away by mother’s blood
○ Birds and other reptiles - egg is surrounded by a shell that is permeable to gases but not to
liquids
■ Any soluble nitrogenous wastes released by embryo would be trapped within the egg and
could accumulate to dangerous levels
■ Hence, uric acid is the waste of reptiles (nontoxic)
 ■ Uric acid is left behind as the animal
hatches
● Amount of nitrogenous waste produced is coupled to:
○ Animal’s energy budget
■ Endotherms - use more energy, eat
more food, produce more nitrogenous
waste vs. ectotherms
○ Diet
■ Predators which derive energy from
protein excrete more nitrogen than
animals that rely mainly on lipids or carbs
as energy sources

EXCRETORY SYSTEMS
● Water balance depends on the regulation of solute
movement between internal fluids and external env.
○ Movement handled by excretory systems
○ These systems are central to homeostasis b/c
they dispose of metabolic wastes and control
body fluid composition

Excretory Processes (key steps)

● Animals produce fluid waste called urine
1.) Filtration
● Body fluid (blood, coelomic fluid, hemolymph) is
brought in contact with selectively permeable
membrane of transport epithelium
● Driven by hydrostatic pressure (blood pressure)
● Cells, proteins, and other large molecules cannot
cross the epithelial membrane and remain in the
body fluid
● Water and small solutes (sugar, salt, amino acids,
nitrogenous wastes) cross the membrane forming a
sol’n called the filtrate
● Filtrate
○ Converted to waste fluid by specific transport
material into or out of the filtrate

2.) Reabsorption
● Selective
● Recovers useful molecules and water from the
filtrate and returns them to the body fluid
● Glucose, certain salts, vitamins, hormones, and amino
acids are reabsorbed by active transport
● Nonessential solutes and wastes are:
○ Left in the filtrate or
○ Added to it by selective secretion

3.) Secretion
● Adds nonessential solutes/wastes to filtrate
 ● Occurs by active transport
● Pumping of various solutes in turn determines whether waste moves by osmosis into or out of the
filtrate

4.) Excretion
● Processed filtrate containing nitrogenous wastes is released from the body as urine

Types of Excretory Systems

1.) Protonephridia
2.) Metanephridia
3.) Malpighian tubules
4.) Kidneys
a.) P
b.) K
c.) K
d.) Mesonephric duct
e.) Metanephric duct

Protonephridia Platyhelminthes ● Consists of a network of dead-end tubules that branch

Flatworms throughout body
(no coelom or body ● Flame bulbs (cellular units)
cavity) ○ Cap the branches of each protonephridium
Some annelids ○ Consists of a tubule cell and cap cell
Mollusc larvae ○ Has a tuft of cilia projecting into tubule
Lancelets

During filtration
● Beating of cilia draws water and solutes from interstitial fluid
through flame bulb
○ Releases filtrate into tubule network
● Processed filtrate moves outward through the tubules and
empties as urine via external openings
● Urine excreted by freshwater flatworms is low in solutes so its
production helps to balance osmotic uptake of water from env.

● Freshwater flatworms
○ Protonephridia serve chiefly in osmoregulation
○ Most metabolic wastes
 ■ Diffuse out of the animal across the body
surface
■ Excreted into gastrovascular cavity and
eliminated through mouth
● Parasitic flatworms
○ Isoosmotic to surround fluids of host organisms have
protonephridia functioning for disposal of nitrogenous
wastes

Metanephridia Annelids ● Excretory organs that collect fluid directly from the
(earthworms) coelom
● Pair of metanephridia found in each segment of annelid
○ Immersed in coelomic fluid and enveloped by
capillary network
● Ciliated funnel surrounds internal opening of each
metanephridium

Process:
● Cilia beat and fluid is drawn into collecting tubule which
includes storage bladder that opens to the outside

● Earthworms inhabit damp soil and usually experience a net

uptake of water by osmosis through their skin
● Metanephridia balance the water influx by producing dilute
urine (hypoosmotic to body fluids)
● Hypoosmotic filtrate allows transport epithelium to
reabsorb most solutes and return them to blood in the
capillaries
● Nitrogenous wastes remain in tubule and excreted to env.
● Metanephridia of earthworm thus serve both excretory and
osmoregulatory function

Malpighian Insects ● Remove nitrogenous wastes and function in

Tubules Other terrestrial osmoregulation
arthropods ● Extend from dead-end tips immersed in hemolymph to
openings into digestive tract

Filtration step is absent, instead:

● Transport epithelium lining the tubules secrete certain
 solutes (+nitro. wastes) from hemolymph into lumen of
tubule
● Water follows solutes into tubule by osmosis
● As fluid passes from tubules into rectum, most solutes are
pumped back into hemolymph
○ Water reabsorption by osmosis follows
● Nitro. Wastes (uric acid) are eliminated as nearly dry matter
along with feces

● Insect excretory system is capable of conserving water

very effectively

Kidneys Vertebrates and ● Compact organ

other chordates ● Functions in both osmoregulation and excretion
● Consists of tubules
○ Arranged in highly organized manner
○ Closely associated with network of capillaries
● Vertebrate excretory system (kidneys)
○ Aksi includes ducts and other structures that carry
urine from tubules out of kidneys and eventually the
body
○ Typically nonsegmented
○ Hagfishes (jawless vertebrates) have kidneys with
segmentally arranged tubules

Human excretory system

● Consists of
○ Kidneys
■ Pair of organs each 10cm in length
○ Organs for transporting urine
○ Organs for storing urine
○ Ureter (two)
 ■ Duct where urine produced in each kidney exits through
■ Drain into urinary bladder (common sac)
■ Urination – urine expelled from bladder through urethra (tube)
○ Urethra empties to the outside
■ Near the vagina – females
■ Through penis – males
■ Sphincter muscles near junction of urethra and bladder – regulate urination

Kidney structure
● Renal Cortex
○ Outer region
○ Supplied with blood by renal artery and drained by renal vein
○ Where tightly packed excretory tubules and associated blood vessels lie
● Renal medulla
○ Inner region
○ Supplied with blood by renal artery and drained by renal vein
○ Where tightly packed excretory tubules and associated blood vessels lie
● Excretory tubules
○ Carry and process a filtrate produced from blood entering kidney
○ Nearly all the fluid in filtrate is reabsorbed into surrounding blood vessels
○ Exit kidney in renal vein
● Renal vein
○ Where reabsorbed fluid from kidney exits through
● Renal Pelvis
○ Remaining fluid (filtrate?) leaves excretory tubules as urine
○ Collected in the inner renal pelvis
○ Exits kidney via ureter
● Ureter
○ Where urine exits through from kidney

Nephrons
● Functional units of vertebrate kidney (cell of kidney)
● Weaves back and forth across renal cortex and renal medulla
1.) Cortical nephrons
● Reach only a short distance into medulla
● 85% of 1 million nephrons are cortical nephrons
2.) Juxtamedullary nephrons
● Extend deep into medulla
 ● Essential for production of urine that is hyperosmotic to body fluids (key adaptation for water
conservation in mammals)

Nephron organization
● Consists of single long tubule and ball of capillaries
● Glomerulus
○ Ball of capillaries
● Bowman’s capsule
○ Cup-shaped swelling
○ Formed by the blind end of the long tubule
○ Surrounds the glomerulus
○ Filtrate is formed when blood pressure
forces fluid from blood in glomerulus into
lumen of Bowman’s capsule
Processing occurs as filtrate passes through 3 major regions of
nephron:
1.) Proximal tubule
2.) Loop of Henle (hairpin turn with descending and
ascending limb)
3.) Distal tubule
● Collecting duct
○ Receives processed filtrate from many nephrons
○ Transports filtrate to renal pelvis
○ Supplied with blood by afferent arteriole
● Afferent arteriole
○ Offshoot of renal artery
○ Branches and forms the capillaries of glomerulus
○ Capillaries converge as they leave the
glomerulus and form an efferent arteriole
● Efferent arteriole
○ Formed by converging capillaries of glomerulus as they exit the glomerulus
○ Branches of efferent arteriole form the peritubular capillaries
■ Surround the proximal and distal tubules
■ Other branches extend downward and form the vasa recta
● Hairpin-shaped capillaries
● Serve the renal medulla (loop of Henle of juxtamedullary nephrons)

Processing of Blood Filtrate

● Human kidney: filtrate forms when fluid passes from bloodstream to lumen of Bowman’s capsule
● Glomerular capillaries and specialized cells of Bowman’s capsule retain blood cells and large
molecules (plasma proteins) but are permeable to water and small solutes
○ Filtrate produced in capsule contains salts, glucose, amino acids, vitamins, nitrogenous
wastes, and other small molecules
○ Concentration of these substances in initial filtrate are the same as those in blood plasma (b/c they
freely pass between glomerular capillaries and capsule)
● Roughly 1,600 L of blood flows through a pair of human kidneys per day
○ 180 L of initial filtrate yielded
○ 99% of water and nearly all sugars, amino acids, vitamins, other organic nutrients are reabsorbed
into blood
○ 1.5 L of urine transported to bladder
Steps from Blood Filtrate to Urine
Glomerulus → Bowman’s capsule → Proximal tubule → descending limb of loop of Henle → Ascending limb of
loop of Henle → Distal tubule → Collecting duct
1.) Proximal tubule
● Reabsorption in proximal tubule is critical for recapture of ions, water, valuable nutrients from huge
volume of initial filtrate
1.) NaCl in filtrate enters cells of transport epithelium by facilitated diffusion and cotransport
mechanisms
2.) NaCl transferred to interstitial fluid by active transport
● This transfer of positive charge out of tubule drives passive transport of Cl-
3.) Water follows by osmosis as salt moves from filtrate to interstitial fluid
● Reduces filtrate volume considerably
4.) Salt and water exiting filtrate diffuse from interstitial fluid into peritubular capillaries
5.) Glucose, amino acids, potassium ions (K-) and other essential substances are also actively or
passively transported from filtrate to interstitial fluid then into peritubular capillaries
● Processing of filtrate in proximal tubule helps maintain a relatively constant pH in body fluids
○ Cells of transport epithelium secrete H- into lumen of tubule and synthesize and secrete
ammonia
○ Ammonia acts as buffer to trap H in form of ammonium ions (NH4+)
■ More acidic filtrate = more ammonia cells produce and secrete
■ Mammal’s urine usually contains some ammonia from this source
○ Proximal tubule also reabsorbs about 90% of buffer bicarbonate (HCO3-) from filtrate
■ Contributed further to pH balance in body fluids
● Materials to be excreted become concentrated as filtrate passes through proximal tubule
○ Many wastes leave body fluids during nonselective filtration process and remain in filtrate
while water and salts are reabsorbed
○ Urea is reabsorbed at a much lower rate than salt and water
○ Materials actively secreted into filtrate from surrounding tissues
○ Drugs and toxins processed in the liver pass from the peritubular capillaries into interstitial fluid
○ Molecules are then actively secreted by transport epithelium into lumen of proximal tubule

2.) Descending limb of loop of Henle

● Filtrate enters loop of Henle upon leaving the proximal tubule
● Further reduces filtrate volume via distinct stages of water and salt movement
1.) Aquaporin proteins (water channels) make transport epithelium freely permeable to water
● Almost no channels for salt and other small solutes (low permeability for these subs.)
2.) Filtrate loses water and increases in solute concentration along its journey down the
descending limb
● Interstitial fluid bathing the tubule is hyperosmotic to the filtrate (hence water moves
out of the tubule by osmosis)
● Osmolarity of interstitial fluid increases progressively from cortex through medulla
3.) Highest osmolarity (1,200 mOsm/L) occurs at the elbow of the loop of Henle

3.) Ascending limb of loop of Henle

1.) Filtrate reaches the tip of the loop and then returns to the cortex in the ascending limb
2.) Has transport epithelium lacking water channels (impermeable to water)
3.) Two specialized regions:
1.) Thin segment
● Near loop tip
 ● As filtrate ascends in thin segment, NaCl (now highly concentrated from descending
limb) diffuses out of permeable tubule into interstitial fluid
● Movement of NaCl out of tubule helps maintain osmolarity of interstitial fluid in
medulla

2.) Thick segment

● Adjacent to distal tubule
● Movement of NaCl out of filtrate continues but:
● Epithelium actively transports NaCl into interstitial fluid
● Filtrate becomes more dilute as it moves up to the cortex in the ascending limb
of loop
● B/c it lost salt, not water

4.) Distal tubule

● Plays a key role in regulating K- and NaCl concentration of body fluids
● Regulation involves variation in amt of K+ secreted into filtrate and the amt of NaCl absorbed from
filtrate
● Also contributes to pH regulation by controlled secretion of H+ and reabsorption of HCO3-

5.) Collecting duct

● Collecting duct processes filtrate into urine
● Carries urine to renal pelvis
1.) As filtrate passes along the transport epithelium of collecting duct, hormonal control of
permeability and transport determines the extent to which the urine becomes concentrated
● When kidneys are conserving water (concentrated urine)
2.) Aquaporin channels in collecting duct allow water molecules to cross epithelium
3.) Epithelium remains impermeable to salt and, in the renal cortex, to urea
4.) Filtrate becomes increasingly concentrated, losing more and more water by osmosis to the
hyperosmotic interstitial fluid
● Happens as collecting duct transverses the gradient osmolarity in kidney
5.) Inner medulla
● Duct becomes permeable to urea
● Some urea diffuses out of the duct and into the interstitial fluid b/c of high urea concentration in
filtrate
● NaCl and Urea contribute to high osmolarity of the interstitial fluid in medulla
● Net result is urine that is hyperosmotic to general body fluids
When producing dilute urine
2.) Collecting duct actively absorbs salts without allowing water to follow by osmosis
3.) Epithelium lacks aquaporin channels
4.) NaCl is actively transported out of filtrate
Solute Gradients and Water Conservation
● Ability of mammalian kidney to conserve water is a key adaptation for terrestrial habitats
● Humans – osmolarity of blood is 300mOsm/L but kidney can excrete urine up to 1,2000mOsm/L
● Mammalian kidney - production of hyperosmotic urine is possible only because considerable
energy is expended for active transport of solutes against concentration gradients
● Nephrons (part. Loop of Henle) can be thought of as energy-consuming machines that produce an
osmolarity gradient suitable for extracting water from the filtrate in the collecting duct
● Primary solutes affecting osmolarity are
○ NaCl - concentrated in the renal medulla by the loop of Henle
○ Urea – passes across epithelium of collecting duct in inner medulla

How the Mammalian Kidney Concentrates Urine

1.) Filtrate passing from Bowman’s capsule to the proximal tubule has abt the same osmolarity as blood
2.) Large amt of water and salt is reabsorbed from the filtrate as it flows through the proximal tubule in the
renal cortex
● So filtrate’s volume decreases substantially but osmolarity stays the same
3.) As filtrate flows from cortex to medulla in the descending limb of loop of Henle
● Water leaves the tubule by osmosis
● Solutes (+ NaCl) become more concentrated and osmolarity of filtrate increases
4.) Diffusion of salt out of the tubule is maximal as filtrate rounds the curve and enters the ascending
limb (permeable to salt but not to water)
● NaCl diffusing from ascending limb helps maintain high osmolarity in interstitial fluid of renal
medulla
 ● Loop of Henle and surrounding capillaries act as a type of countercurrent system to
generate the steep osmotic gradient between the medulla and cortex
● Use of countercurrent exchange:
○ Some endotherms have countercurrent heat exchange to reduce heat loss (passive)
○ Countercurrent gas exchange in fish gills maximize oxygen absorption (passive)
● Countercurrent Multiplier Systems
○ Countercurrent system in kidney; NOT PASSIVE
○ Countercurrent system of loop of Henle involves active transport and thus expend
energy
○ The active transport of NaCl from filtrate in upper part of ascending limb of loop of Henle
maintains a high salt concentration in the interior of the kidney
○ Enables kidney to form concentrated urine
○ Descending and ascending vessels of vasa recta carry blood in opposite directions
through kidney’s osmolarity gradient
○ As descending vessel conveys blood toward inner medulla, water is lost from blood and
NaCl gained by diffusion
○ Reversed net fluxes as blood flows back toward cortex in ascending vessel of vasa recta
with water reentering blood and salt diffusing out
○ Vasa recta can supply the kidney with nutrients and other important substances carried
by blood without interfering with osmolarity gradient in inner and outer medulla
○ Countercurrent-like characteristics of loop of Henle and vasa recta help to generate steep
osmotic gradient bet. Medulla and cortex
○ Thick segment of ascending limb of loop of Henle is where expenditure of energy to
maintain osmotic gradient occurs
■ Also where NaCl is actively transported out of tubule
○ Filtrate is actually hypoosmotic to body fluids by the time it reaches the distal tubule
5.) Filtrate descends again toward to medulla (in collecting duct) where it’s permeable to water but not to salt
● Osmosis extracts water from filtrate as it passes from cortex to medulla and encounters
interstitial fluid of increasing osmolarity
● This process concentrates salt, urea, and other solutes in filtrate
● Some urea passes out of lower portion of collecting duct and contributes to high interstitial
osmolarity of inner medulla
6.) Urine reaches 1,200 mOsm/L (same as osmolarity of interstitial fluid in inner medulla) when kidney
concentrates urine maximally
● Urine is hyperosmotic to blood and interstitial fluid elsewhere in body but is isoosmotic with
inner medulla’s interstitial fluid
● This high osmolarity allows solutes remaining in urine to be excreted from body with minimal water
loss
Adaptations of Vertebrate Kidney
● Presence of juxtamedullary nephrons is a kedy adaptation the enables terrestrial mammals to shed
salts and nitrogenous wastes without squandering water
● Differences in length of loop of Henle in juxtamedullary nephrons and in no. Of juxtamedullary and cortical
nephrons help to fine-tune osmoregulation to particular habitats

1.) Mammals
○ Desert mammals (Australian hopping mice, North American kangaroo rats, etc.)
■ Excrete most hyperosmotic urine
■ Have many juxtamedullary nephrons with loops of Henle that extend deep into medulla
■ Long loops maintain steep osmotic gradients in kidney, resulting to urine becoming
very concentrated as it passes from cortex to medulla in collecting ducts
○ Aquatic mammals (beavers, muskrats, etc.) in fresh water
■ Have mostly cortical nephrons
■ Lower ability to concentrate urine
○ Terrestrial mammals (in moist conditions)
■ Have loops of Henle intermediate in length
■ Intermediate capacity to produce urine intermediate in concentration to that of freshwater
and desert mammals
2.) Vampire Bat
○ Feeds at night on blood of large birds and mammals
○ Uses sharp teeth to make a small incision in pey’s skin and laps up blood from wound
○ Anticoagulants in bat’s saliva prevent blood from clotting
○ Kidneys excrete large volumes of dilute urine
■ This is so the bat could lose weight to take off
○ Vampire bat faces another problem in the roost (cave)
○ Nutrition it derives from blood comes in the form of protein
○ Digesting proteins generates large quantities of urea but roosting bats lack access to drinking
water necessary to dilute it
○ Solution: kidneys shift to producing small quantities of highly concentrated urine
 ■ Disposes urea load while conserving water
○ Vampire bat’s ability to alternate rapidly between producing large amts of dilute urine and
small amts of very hyperosmotic urine is an essential part of its adaptation to an unusual food
source
3.) Birds and other Reptiles
● Birds usually live in dehydrating envs.
○ Birds have juxtamedullary nephrons like mammals
○ Nephrons of birds have loops of Henle that extend less far into medulla than those of mammals
○ Bird kidneys cannot concentrate urine to high osmolarities achieved by mammalian kidneys
○ Birds can produce hyperosmotic urine, but their main water conservation adaptation is having
uric acid as nitrogenous waste
● Other reptiles only have cortical nephrons
○ Produce urine that is isoosmotic or hypoosmotic to body fluids
○ Epithelium of cloaca from which urine and feces leave the body conserves fluid by reabsorbing
water from these wastes
○ Also excrete nitrogenous wastes as uric acid
4.) Freshwater Fishes and Amphibians
● Freshwater fishes
○ Hyperosmotic to their surroundings
○ Produce large volumes of very dilute urine
○ Kidneys are packed with cortical nephrons
○ Produce filtrate at high rate
○ Salt conservation relies on reabsorption of ions from the filtrate in distal tubules
● Amphibians
○ Function like those of freshwater fishes when in freshwater
○ Kidneys excrete dilute urine while their skin accumulates certain salts from the water by active
transport
○ On land (dehydration problem of osmoreg.)
○ Frogs conserve body fluid by reabsorbing water across the epithelium of urinary bladder
5.) Marine Bony Fishes
● Fewer and smaller nephrons
● Nephrons lack a distal tubule
● Kidneys have small glomeruli or lack glomeruli entirely
● Thus, filtration rates are low and very little urine is excreted
● Main function of kidney in marine bony fishes:
○ Get rid of divalent ions (Ca2+, Mg2+, SO42-)
○ They take in these ions by incessantly drinking seawater
○ Rid themselves of these ions by secreting them into proximal tubules of nephrons and
excreting them in urine
○ Osmoregulation in marine bony fishes also relies on specialized chloride cells in gills
○ Chloride cells maintain proper levels of monovalent ions (Na+ and Cl-)
○ Generation of ion gradients and movement of ions across membranes are central to salt and water
balance in marine bony fishes

Hormonal Circuits Link Kidney function, Water Balance, and Blood Pressure
● In mammals, both volume and osmolarity of urine are adjusted acc. To banimal’s water and salt balance
and its rate of urea production
● High salt intake and low water = urea excreted in small volumes of hyperosmotic urine (minimal
water loss)
 ● Scarce salt and high water = kidney eliminates excess water with little salt loss by excreting
hypoosmotic urine (dilute)
Homeostatic Regulation of Kidney
● Combination of nervous and hormonal control manages the osmoregulatory function of mammalian
kidney

Antidiuretic Hormone (ADH)

● Key hormone in of the kidney
● Released from posterior pituitary
● Bind to and activate membrane receptors on surface of
collecting duct cells
○ Activated receptors initiate signal transduction
cascade that directs insertion of aquaporin
proteins into membrane lining collecting duct
● More aquaporin channels result in more water recapture,
reducing urine volume (high level of urine production is
called dieresis; ADH is thus antidiuretic hormone)

What occurs when blood osmolarity rises (after eating salty

food or losing water through sweating)
● Osmolarity rises above normal range (285-295
mOsm/L)
○ Osmoreceptor cells in hypothalamus trigger
increased release of ADH from posterior
pituitary
○ Resulting increase in water reabsorption in
collecting duct
■ Concentrates urine
■ Reduces urine volume
■ Lowers blood osmolarity
back toward setpoint
○ As osmolarity in blood falls
■ Negative feedback
mechanism reduces
activity of osmoreceptor
cells in hypothalamus
■ ADH secretion reduced
You drink lots of water and don’t sweat profusely
● Blood osmolarity falls below set point
● Drop in ADH secretion to a very low
level
● Resulting decrease in permeability of
collecting ducts
○ Reduce water reabsorption
○ Large dilute urine volume

● Blood osmolarity, ADH release, and water reabsorption in the kidney are normally linked in a feedback
circuit that contributes to homeostasis
○ Disrupting this circuit can interfere with water balance
○ E.g. alcohol inhibit ADH release - leads to excessive urinary water loss/dehydration
Renin-Angiotensin-Aldosterone System (RAAS)
● Release of ADH is a response to an increase in blood osmolarity, when body is dehydrated from
excessive water loss or inadequate water intake
● Excessing loss of both water and salt (e.g. with diarrhea) will reduce blood volume without increasing
osmolarity
○ Will not affect ADH release
but something else
● Endocrine circuit, renin-
angiotensis-aldosterone system
(RAAS) also regulates kidney
function
● Regulates blood volume and
pressure
● RAAS
○ Responds to drop in blood
volume and pressure by
increasing water and Na+
reabsorption
○ Involves the juxtaglomerular
apparatus (JGA)
■ Specialized tissue
consisting of cells of and around the afferent arteriole, which supplies blood to glomerulus
■ When blood pressure or volume drops in afferent arteriole (e.g. b/c of dehydration) the JGA
releases the enzyme Renin
○ Renin
■ Initiates a sequence of steps that cleave a plasma protein, angiotensinogen
● Yields a peptide, angiotensin II
○ Angiotensin II
■ Hormone
■ Triggers vasoconstriction
● Increases BP and decreases blood flow to capillaries in kidney
■ Stimulates the adrenal glands to release aldosterone
○ Aldosterone
■ Causes nephrons’ distal tubules and collecting duct to reabsorb more Na- and water
● Increases BP and blood volume
○ Drugs that block angiotensin II production are widely used to treat hypertension
■ B/c it results in increased BP
■ These drugs are specific inhibitors of angiotensin converting enzyme (ACE) which
catalyzes one of the steps in the production of angiotensin II
● RAAS operates as a feedback circuit
○ Drop in BP and blood volume triggers renin release
○ Resulting production of angiotensin II and release of aldosterone cause a rise in BP and volume
○ This reduces the release of renin from JGA
● Both ADH and RAAS increase water reabsorption in the kidney
○ ADH alone would lower blood Na+ concentration via water reabsorption in kidney
○ RAAS helps maintain body fluid osmolarity at set point by stimulating Na+ reabsorption

Atrial natriuretic peptide (ANP)

● Opposes RAAS
 ● Walls of atria of heart release ANP in response to increase in BP and volume
● ANP inhibits the release of renin from JGA
● ANP inhibits NaCl reabsorption by collecting ducts
● ANP reduces aldosterone release from adrenal glands
● These all lower BP and volume
○ Thus, ADH, RAAS, and ANP all regulate kidney’s ability to control blood osmolarity, salt
concentration, volume, and pressure (check and balances)

Thirst
● Plays an essential role in control of water and salt balance
● Some neurons in hypothalamus are dedicated in regulating thirst
● Stimulating one set of neurons in mice causes intense drinking behavior (even if hydrated)
● Stimulating a second set causes immediate halt in water consumption (even if dehydrated)