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    Chapter 44

    Osmoregulation and

    ExcretionRec. Dur.: 4h (8-9)

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    Gross structure of the kidney

    Detailed structure of the nephron and associatedblood vessels

    Osmoregulation

    Excretion

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    At the end of the lesson, u should be able to:

    A. Anatomy of Mammalian kidney

    Describe how the processes of osmoregulation and excretioncontribute to fluid and electrolyte homeostasis.

    Outline the gross structure of the kidney and the detailedstructure of the nephron and associated blood vessels

    with emphasis on transport processes and their relevant

    proteins (e.g., transport of glucose, Na+, Cl-, water, K+).

    Ultrafiltration

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    At the end of the lesson, u should be able to:

    A. Anatomy of Mammalian kidney

    Trace a drop of filtrate from Bowmans capsule to its releasefrom the body as urine

    Interpret the histology of the kidney, as seen in sections underthe light microscope.

    Explain the functioning of the kidney in the control of water andmetabolic waste levels of the body fluids.

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    At the end of the lesson, u should be able to:

    B. Renal processes of regulation of water and solutes

    Describe the hormonal regulation of fluid and electrolytebalance by antidiuretic hormone (ADH) and atrial natriureticpeptide (ANP).

    KIV 2011?: Outline the mechanism of dialysis in the case of

    kidney failure.

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    Overview: A Balancing Act

    Osmoregulation regulates soluteconcentrations and balances the gain and lossof water

    Excretion gets rid of nitrogenous metabolitesand other waste products

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    Fig. 44-2

    Selectively permeablemembrane

    Net water flow

    Hyperosmotic side Hypoosmotic side

    Water

    Solutes

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    Osmosis and Osmolarity

    Cells require a balance between osmotic gainand loss of water

    Osmolarity, the solute concentration of a

    solution, determines the movement of wateracross a selectively permeable membrane

    If two solutions are isoosmotic, the movement

    of water is equal in both directions If two solutions differ in osmolarity, the net flow

    of water is from the hypoosmotic to the

    hyperosmotic solution Adequate self-study material

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    Urea

    The liver of mammals and most adultamphibians converts ammonia to less toxicurea

    The circulatory system carries urea to thekidneys, where it is excreted

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    Concept 44.3: Diverse excretory systems arevariations on a tubular theme

    Excretory systems regulate solute movementbetween internal fluids and the externalenvironment

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    Fig. 44-14a

    A:

    B: (Red) & (blue)

    E:

    D:

    C:

    F:

    (a) Excretory organs and major

    associated blood vessels

    G:

    Can u name these?

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    Fig. 44-14b

    (b) Kidney structureSection of kidneyfrom a rat 4 mm

    C:

    B:

    D:

    A:

    Can u name these?

    Fi 44 14

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    Fig. 44-14cB:A:

    E:

    (c) Nephron types

    Torenalpelvis

    D:

    C:Can u name these?

    Fi 44 14d

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    Fig. 44-14d

    A:

    B:

    SEM

    C:

    E:

    F:D:

    (d) Filtrate and blood flow

    G:

    H:

    i:

    J:

    K:

    L:M:

    10 m

    Can u namethese?

    Fi 44 10

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    Fig. 44-10

    Capillary

    Excretion

    Secretion

    Reabsorption

    Excretorytubule

    Filtration

    Urine

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    Excretory Processes

    Most excretory systems produce urine byrefining a filtrate derived from body fluids

    Key functions of most excretory systems:

    Filtration: pressure-filtering of body fluids

    Reabsorption: reclaiming valuable solutes

    Secretion: adding toxins and other solutesfrom the body fluids to the filtrate

    Excretion: removing the filtrate from the

    system Adequate self-study material

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    Kidneys

    Kidneys, the excretory organs of vertebrates,function in both excretion and osmoregulation

    Adequate self-study material

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    Structure of the Mammalian Excretory System

    The mammalian excretory system centers onpaired kidneys, which are also the principal siteof water balance and salt regulation

    Each kidney is supplied with blood by a renalartery and drained by a renal vein

    Urine exits each kidney through a duct calledthe ureter

    Both ureters drain into a common urinarybladder, and urine is expelled through aurethra

    Adequate self-study material

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    The mammalian kidney has two distinctregions: an outer renal cortex and an innerrenal medulla

    Adequate self-study material

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    The nephron, the functional unit of thevertebrate kidney, consists of a single longtubule and a ball of capillaries called theglomerulus

    Bowmans capsule surrounds and receivesfiltrate from the glomerulus

    Adequate self-study material

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    Filtration of the Blood

    Filtration occurs as blood pressure forces fluidfrom the blood in the glomerulus into the lumenof Bowmans capsule

    Filtration of small molecules is nonselective

    The filtrate contains salts, glucose, aminoacids, vitamins, nitrogenous wastes, and other

    small molecules

    Adequate self-study material

    P h f h Fil

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    Pathway of the Filtrate

    From Bowmans capsule, the filtrate passesthrough three regions of the nephron: theproximal tubule, the loop of Henle, and thedistal tubule

    Fluid from several nephrons flows into acollecting duct, all of which lead to the renalpelvis,which is drained by the ureter

    Cortical nephrons are confined to the renalcortex, while juxtamedullary nephrons haveloops of Henle that descend into the renalmedulla

    Adequate self-study material

    Bl d V l A i d i h h N h

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    Blood Vessels Associated with the Nephrons

    Each nephron is supplied with blood by anafferent arteriole, a branch of the renal arterythat divides into the capillaries

    The capillaries converge as they leave theglomerulus, forming an efferent arteriole

    The vessels divide again, forming the

    peritubular capillaries, which surround theproximal and distal tubules

    Adequate self-study material

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    Vasa recta are capillaries that serve the loopof Henle

    Adequate self-study material

    C t 44 4 Th h i i d f

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    Concept 44.4: The nephron is organized forstepwise processing of blood filtrate

    The mammalian kidney conserves water byproducing urine that is much moreconcentrated than body fluids

    Adequate self-study material

    LE 26-12

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    Nephron loop

    Urine storage

    and elimination

    Reabsorption

    Secretion

    Glomerular capsule

    KEY

    Glomerulus

    Proximal convoluted tubule Distal convolutedtubule

    Collecting duct

    Leak channel

    Countertransport

    Exchange pump

    Cotransport

    Diffusion

    Tubular fluid

    Cells ofproximal

    convolutedtubule

    Glucose andother organic

    solutes

    H+ HCO3 CO2 H2OH2CO3

    H+

    H+

    Na+ Na+Na+

    HCO3

    CO2

    H2CO3

    H+

    H2O

    HCO3

    Na+

    H+

    Facilitateddiffusion

    HCO3

    Na+

    2 K+

    3 Na+

    2 K+

    3 Na+

    Peritubularcapillary

    FYI only

    LE 26-12-3

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    Reabsorption

    Secretion

    KEYLeak channel

    Countertransport

    Exchange pump

    Cotransport

    Diffusion

    Tubular fluid

    Cells ofproximal

    convoluted

    tubule

    Glucose andother organic

    solutes

    H+ HCO3 CO2 H2OH2CO3

    H+

    H+

    Na+ Na+Na+

    HCO3

    CO2

    H2CO3

    H+

    H2O

    HCO3

    Na+

    H+

    Facilitateddiffusion

    Na+

    2 K+

    3 Na+

    Peritubularcapillary

    HCO3

    3 Na+

    2 K+

    FYI only

    F Bl d Filt t t U i A Cl L k

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    From Blood Filtrate to Urine: A Closer Look

    Proximal Tubule

    Reabsorption of ions, water, and nutrientstakes place in the proximal tubule

    Molecules are transported actively andpassively from the filtrate into the interstitialfluid and then capillaries

    Some toxic materials are secreted into thefiltrate

    The filtrate volume decreases

    Adequate self-study material

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    The function of the loop of

    Henle is to concentrate sodiumions and chloride ions in the

    interstitial fluid in the medulla.

    Play animation

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    Descending Limb of the Loop of Henle Reabsorption of water continues through

    channels formed by aquaporin proteins

    Movement is driven by the high osmolarity ofthe interstitial fluid, which is hyperosmotic tothe filtrate

    The filtrate becomes increasingly concentrated

    Adequate self-study material

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    The epithelium of the ascending limb is permeable to sodiumions and chloride ions but not to water.

    As filtrate ascends in the thin segment of the ascending limb,

    sodium ions and chloride ions diffuse into the interstitial fluid.

    Play animation

    LE 26-13P i l l t d t b l KEYT b l fl id

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    Nephron loop

    Urine storageand elimination

    Glomerular capsule

    Glomerulus

    Proximal convoluted tubuleDistal convoluted

    tubule

    Collecting duct

    Reabsorption

    Secretion

    KEY

    Leak channel

    Exchange pump

    Cotransport

    Diffusion

    Na+CI

    H2O

    2 CI

    Na+K+

    Tubular fluid

    PeritubularfluidNa+

    K+K+

    K+

    CI

    K+

    CORTEX

    DCT andcollecting duct(variablepermeabilityto water;impermeable

    to urea)

    Papillary duct(permeabilityto urea)

    Thin descendinglimb (permeable

    to water,impermeable

    to urea)

    MEDULLA Urea120012001200

    900900

    600300

    300

    300

    100

    H2O

    H2O

    H2O

    H2O

    H2O

    1200

    900

    600

    300 100

    H2O

    300

    H2O

    600

    900

    Na+CI

    Na+CI Na+CI

    Na+CI

    Na+CI

    300mOsm/L

    1200

    Thindescendinglimb(permeableto water;impermeableto solutes)

    Thickascending

    limb(impermeable

    to water;active solute

    transport)

    The mechanism of sodium and chloride ion transportinvolves the Na+K+/2CIcarrier at the apical surfaceand two carriers at the basal surface of the tubularcell: a potassiumchloride cotransport pump anda sodiumpotassium exchange pump. The net resultis the transport of sodium and chloride ions into theperitubular fluid.

    The permeability characteristics of both the loop and the collectingduct tend to concentrate urea in the tubular fluid and in the medulla.The nephron loop, DCT, and collecting duct are impermeable to urea.As water reabsorption occurs, the urea concentration rises. Thepapillary ducts permeability to urea accounts for roughly one -third

    of the solutes in the deepest portions of the medulla.

    Active transport of NaCI along the ascendingthick limb results in the movement of water fromthe descending limb.

    FYI only

    LE 26-13a

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    Reabsorption

    Secretion

    KEY

    Leak channel

    Exchange pump

    Cotransport

    Diffusion

    2 CI

    Na+

    Tubular fluid

    Peritubularfluid

    K+

    K

    +

    CI

    Na+

    K+

    K+K+

    FYI only

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    Ascending Limb of the Loop of Henle In the ascending limb of the loop of Henle, salt

    but not water is able to diffuse from the tubule

    into the interstitial fluid The filtrate becomes increasingly dilute

    Adequate self-study material

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    Water is able to move out by osmosis because ofthe high osmotic concentration of the interstitial fluid

    in the medulla.

    Play anima

    Play animation

    LE 26-14

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    Nephron loop

    Urine storageand elimination

    Glomerularcapsule

    Glomerulus

    Proximalconvoluted tubule

    Distal convoluted tubule

    Collecting duct

    Reabsorption

    Secretion

    KEY

    Leak channel

    Exchange pump

    Cotransport

    Diffusion

    Countertransport

    CINa+

    Tubular fluid

    K+

    Entire DCT

    Na+

    Na+K+

    Na+K+

    Na+K+

    Na+

    K+

    Na+

    K+

    Na+

    CI

    Na+CI

    Peritubularcapillary

    Sodium and chloride

    reabsorption

    Aldosterone-sensitiveportion of DCT and

    collecting duct

    Sodiumpotassium exchange

    FYI only

    LE 26-14aTubular fluid

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    Reabsorption

    Secretion

    KEY

    Leak channel

    Exchange pump

    Cotransport

    Diffusion

    Countertransport

    CINa+

    Tubular fluid

    K+

    Entire DCT

    Na+

    Na+

    K+

    K+

    CI

    Na+CI

    Peritubularcapillary

    Sodium and chloride

    reabsorption

    FYI only

    LE 26-14b

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    Reabsorption

    Secretion

    KEY

    Leak channel

    Exchange pump

    Cotransport

    Diffusion

    Countertransport

    Na+

    Na+K+

    Na+K+

    Na+K+

    Na+

    Aldosterone-sensitiveportion of DCT and

    collecting duct

    Sodiumpotassium exchange

    FYI only

    LE 26-14cTubular Hydrochloric Ammonium

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    Reabsorption

    Secretion

    KEY

    Leak channel

    Exchange pump

    Cotransport

    Diffusion

    Countertransport

    H+

    Entire DCTand

    collectingduct

    H+

    secretion and HCO3

    reabsorption

    HCO3 Sodium bicarbonate

    Peritubularcapillary

    CO2

    Na+ HCO3

    H+HCO3

    CO2

    Na+

    HCO3

    CI

    H+HCO3

    H+

    HCO3

    H+

    H+

    CO2

    CO2

    CI

    CI

    CI

    Na+

    Na+

    Na+

    HCO3

    HCO3

    CI

    CI

    CI

    H2CO3

    H2O

    NH4

    NH4

    NH4

    Aminoacid

    deamination

    fluidy

    acid chloride

    FYI only

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    Distal Tubule

    The distal tubule regulates the K+ and NaClconcentrations of body fluids

    The controlled movement of ions contributes topH regulation

    Adequate self-study material

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    Collecting Duct

    The collecting duct carries filtrate through themedulla to the renal pelvis

    Water is lost as well as some salt and urea,and the filtrate becomes more concentrated

    Urine is hyperosmotic to body fluids

    Adequate self-study material

    Fig. 44-15

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    Key

    Activetransport

    Passivetransport

    INNERMEDULLA

    OUTERMEDULLA

    H2O

    CORTEX

    Filtrate

    Loop ofHenle

    H2O K+HCO3

    H+ NH3

    Proximal tubule

    NaCl Nutrients

    Distal tubule

    K+ H+

    HCO3

    H2O

    H2O

    NaCl

    NaCl

    NaCl

    NaCl

    Urea

    Collectingduct

    NaCl

    By what process are

    substances moving?

    Fig. 44-16-1

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    Key

    Activetransport

    Passivetransport

    INNERMEDULLA

    OUTERMEDULLA

    CORTEXH2O

    300300

    300

    H2O

    H2O

    H2O

    400

    600

    900

    H2O

    H2O

    1,200

    H2O

    300

    Osmolarity ofinterstitial

    fluid(mOsm/L)

    400

    600

    900

    1,200

    In which direction ismedullary/ interstitialfluid concentration

    gradient?By what process isH2O moving?What causes it tomove?

    Effects oninterstitial fluid?Is this consideredreabsorption or

    excretion?

    Fig. 44-16-2

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    Key

    Activetransport

    Passivetransport

    INNERMEDULLA

    OUTERMEDULLA

    CORTEXH2O

    300300

    300

    H2O

    H2O

    H2O

    400

    600

    900

    H2O

    H2O

    1,200

    H2O

    300

    Osmolarity ofinterstitial

    fluid(mOsm/L)

    400

    600

    900

    1,200

    100

    NaCl

    100

    NaCl

    NaCl

    NaCl

    NaCl

    NaCl

    NaCl

    200

    400

    700

    By what process areNa+, Cl- moving?What causes them to

    move?Effects on interstitialfluid?Is this consideredreabsorption or

    excretion?

    Fig. 44-16-3

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    Key

    Activetransport

    Passivetransport

    INNERMEDULLA

    OUTERMEDULLA

    CORTEXH2O

    300300

    300

    H2O

    H2O

    H2O

    400

    600

    900

    H2O

    H2O

    1,200

    H2O

    300

    Osmolarity ofinterstitial

    fluid(mOsm/L)

    400

    600

    900

    1,200

    100

    NaCl

    100

    NaCl

    NaCl

    NaCl

    NaCl

    NaCl

    NaCl

    200

    400

    700

    1,200

    300

    400

    600

    H2O

    H2O

    H2O

    H2O

    H2O

    H2O

    H2O

    NaCl

    NaCl

    Urea

    Urea

    Urea

    H2ONa+, Cl-UreaBy what process

    are these moving?What causes it tomove?Effects on interstitialfluid?Is this consideredreabsorption orexcretion?

    Solute Gradients and Water Conservation

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    Solute Gradients and Water Conservation

    Urine is much more concentrated than blood

    The cooperative action and precisearrangement of the loops of Henle and

    collecting ducts are largely responsible for theosmotic gradient that concentrates the urine

    NaCl and urea contribute to the osmolarity of

    the interstitial fluid, which causes reabsorptionof water in the kidney and concentrates theurine

    Adequate self-study material

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    In the proximal tubule, filtrate volumedecreases, but its osmolarity remains the same

    Adequate self-study material

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    The collecting duct conducts filtrate through theosmolarity gradient, and more water exits thefiltrate by osmosis

    Urea diffuses out of the collecting duct as ittraverses the inner medulla

    Urea and NaCl form the osmotic gradient that

    enables the kidney to produce urine that ishyperosmotic to the blood

    Adequate self-study material

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    Animals that musttightly regulate body

    water stores (desertanimals) have verylong loops of Henleand large osmoticgradients

    Brooker1e

    Mammals

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    Mammals

    The juxtamedullary nephron contributes towater conservation in terrestrial animals

    Mammals that inhabit dry environments have

    long loops of Henle, while those in fresh waterhave relatively short loops

    Adequate self-study material

    Dehydration

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    Playanimation

    Can u.c.d. aquaporin?

    Perspiration causes loss of water and sodium ions.

    Blood levels of ADH and aldosterone increase

    Aldosterone increases no. of Na+/K+ pumps (yellow) & Na+,

    K+ channels Body fluid is conserved and urine volume decreases.

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    Playanimation

    Dehydration

    In severe dehydration condition,

    low volume of urine excreted

    may be concentrated to about 1400 mOsm/L

    Adequate self-study material after viewing anime:Overhydration

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    Play

    animation

    Slurrrrpp

    Thirsting for

    exam tips?

    Can u.c. any

    aquaporin?

    Overhydration triggers a decrease in ADH and aldosterone

    No. of Na+/K+ pumps & Na+, K+ channels decreases

    Reabsorption of water and sodium ions decreasesUrine volume increases

    Adequate self-study material after viewing anime:Overhydration

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    Playanimation

    ADH levels are very low or absent

    Duct cells remain impermeable to water and urea

    Urine is very dilute and high in volume Final urine may have an osmolarity as low as 100 mOsm/L

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    Playanimation

    Adequate self-study material after viewing anime:With normal

    hydration, normal ADH results in a few water channels ADH facilitates the diffusion of urea out of the medullary

    collecting duct

    Urea is responsible for up to 40% of the interstitial fluid

    osmolarity Urea circulates back into the loop of Henle and returns to

    collecting ducts

    Urine is concentrated to about twice normal body osmolarity

    (600 mOsm/L)

    Fox 11e: Table 17.2

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    Summary of transport processes in renal tubule &collecting duct

    Adequate self-study material

    Antidiuretic Hormone

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    The osmolarity of the urine is regulated bynervous and hormonal control of water and saltreabsorption in the kidneys

    Antidiuretic hormone (ADH) increases waterreabsorption in the distal tubules and collectingducts of the kidney

    An increase in osmolarity triggers the releaseof ADH, which helps to conserve water

    Adequate self-study material

    Fig. 44-19

    Osmoreceptors inINTERSTITIALFLUID

    COLLECTINGDUCT

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    Thirst

    Drinking reducesblood osmolarity

    to set point.

    phypothalamus trigger

    release of ADH.

    Increasedpermeability

    Pituitarygland

    ADH

    Hypothalamus

    Distaltubule

    H2O reab-sorption helpsprevent further

    osmolarityincrease.

    STIMULUS:Increase in blood

    osmolarity

    Collecting duct

    Homeostasis:Blood osmolarity

    (300 mOsm/L)

    (a)

    Exocytosis

    (b)

    Aquaporinwaterchannels

    H2O

    H2O

    Storagevesicle

    Second messengersignaling molecule

    cAMP

    FLUID

    ADH

    receptor

    ADH

    DUCTLUMEN

    COLLECTINGDUCT CELL

    *nervous and hormonalcontrols involved in theregulation of kidney

    function

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    Mutation in ADH production causes severedehydration and results in diabetes insipidus

    Alcohol is a diuretic as it inhibits the release of

    ADH

    Adequate self-study material

    Fig. 44-21-3 Liver*Why does

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    Renin

    Distaltubule

    Juxtaglomerularapparatus (JGA)

    STIMULUS:Low blood volumeor blood pressure

    Homeostasis:Blood pressure,

    volume

    Angiotensinogen

    Angiotensin I

    ACE

    Angiotensin II

    Adrenal gland

    Aldosterone

    Arterioleconstriction

    Increased Na+and H2O reab-

    sorption indistal tubules

    aldosterone,increase

    blood volumeandpressure?

    Na+ causes:

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    heart to make more ANP Atrial natriuretic peptide

    (ANP) secreted from atriaof heart to decrease sodiumreabsorption in kidney

    *What does this to bloodconcentration of Na+?

    Brooker1e (text & pix)

    The Renin-Angiotensin-Aldosterone System

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    The renin-angiotensin-aldosterone system(RAAS) is part of a complex feedback circuitthat functions in homeostasis

    A drop in blood pressure near the glomeruluscauses the juxtaglomerular apparatus (JGA)to release the enzyme renin

    Renin triggers the formation of the peptideangiotensin II

    Adequate self-study material

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    Angiotensin II

    Raises blood pressure and decreases bloodflow to the kidneys

    Stimulates the release of the hormonealdosterone, which increases blood volumeand pressure

    Adequate self-study material

    Homeostatic Regulation of the Kidney

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    ADH and RAAS both increase waterreabsorption, but only RAAS will respond to adecrease in blood volume

    Another hormone, atrial natriuretic peptide(ANP), opposes the RAAS

    ANP is released in response to an increase in

    blood volume and pressure and inhibits therelease of renin

    Adequate self-study material

    You should now be able to:

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    1. Define osmoregulation, excretion

    2. Using a diagram, identify and describe thefunction of each region of the nephron

    3. Explain how the loop of Henle enhanceswater conservation

    4. Describe the nervous and hormonal controls

    involved in the regulation of kidney function

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    THE END