18 - Excretion System

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    Excretory SystemKK Chapter 14, Hildebrand Chapter 15

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    Excretory System Functions

    1) Maintain salt/water balance by releasing controlledamounts of both under the influence of the endocrinesystem (adrenal and pituitary glands).

    2) Eliminate toxic wastes, especially of protein metabolism.De-amination of amino acids leads to NH4

    +

    (ammonium). This is readily excreted from the gills inmost fish as NH4,

    + but otherwise the liver may turn NH4+

    into less toxic urea. In birds and reptiles, the kidneysturn NH4

    + into uric acid.

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    The Nephron - the basic unit of the kidney

    Renal corpuscle = renal (Bowmans) capsule and glomerulus

    Nephron = renal corpuscle + tubule

    KK 14.2, H&G 15.5

    The renal corpuscle is

    the filter of the kidney,

    where filtrate passesfrom the blood-vascular

    system to the capsule,

    while the tubule

    reclaims solutes from

    the filtrate and conductsurine towards the

    outside of the body.

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    Development of the Kidney

    The kidney tissue develops from thethe mesomere, which swells to create

    a nephric ridge on the dorsal wall of

    the coelom. It develops from front to

    back, changing morphology as it does

    so. The first segments of themesomere produce one pair of

    glomeruli and nephrons per segment,

    and this anterior first kidney is called

    the pronephros. A branch of the

    dorsal aorta in each segment leads to aglomerulus, and each is associated

    with a capsule and tubule.

    KK 14.3a, H&G 15.1

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    Development of the Kidney 2KK 14.3b&c, H&G 15.1

    The pronephros may have originally emptied

    into the coelom, as it does in hagfishes, but

    otherwise collects in a nephric duct that

    transports the urine towards the cloaca. The

    pronephros is a functional kidney in larval

    cyclostomes, many fishes and Amphibia. Inamniotes, even in their embryonic stages, the

    pronephros appears but is never a functional

    kidney.

    In teleosts, the pronephros

    may persist and have a

    glandular role. Physiologists

    refer to it as head kidney.KK 14.3

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    Vertebrate kidneys develop from from the

    mesomere, front to back, beginning with

    the pronephros. Behind the pronephros, a

    second kind of kidney, the mesonephros,

    develops. It has many corpuscles per

    segment. It becomes the adult kidney innon-amniotes, and is then called the

    opisthonephros(tail kidney).

    In amniotes, a third kidney or

    metanephros develops behind theembryonic mesonephros. It does not

    share the archinephric duct but rather a

    new duct emerges from the cloaca to meet

    it. In the adult, this duct is called the

    ureter rather than a metanephric duct.

    Development

    of the Kidney 3KK 14.4, H&G 15.2

    KK 14.4

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    Amniote embryo illustrating all 3 kidney types

    KK 14.5, H&G 15.2

    Note that the same

    duct can be called the

    pronephric duct, the

    mesonephric duct,

    opisthonephric duct,the archinephric duct,

    or simply the nephric

    duct. And, as weshall see, thats not all!

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    Opisthonephric Kidney of a (Male) Salamander

    KK 14.8

    There are many renalcorpuscles per segment.

    The anterior portion of

    the male kidney is

    intimately associated

    with the testis, and acts

    as a sperm storage

    organ. They share the

    same duct leading to

    the cloaca.

    KK 14.8

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    Mammalian Kidney FunctionThe blood supply to the

    glomerulus is from the renalartery. After leaving the

    glomerulus, the blood travels

    to more capillaries around the

    nephric tubule that re-gain

    needed water and solutes lost

    to the capsule. If a renal

    portal vein is present, it

    contributes to these

    capillaries.

    Na+ ions are activelyremoved, and Cl- water

    follows passively.

    KK 14.14, H&G 15.8

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    Mammalian Kidney Function 2The intermediate part of the tubule is a

    site of active tranport of Na+

    into thesurronding tissue, resulting in a very high

    Na+ concentration in that tissue. Cl-

    follows Na+ to maintain charge balance.

    The high salt concentration in the tissue

    causes much of the water still in the urine

    to be reclaimed as it passes down the

    collecting tubule.

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    Human Metanephric KidneyThe human kidneys contain 2 x106 nephrons, whose capsules have 0.76 m2 of filter

    area, or about 1/2 the surface area of the body. There is about 120 km of tubule.

    They receive about 20% of the cardiac output, so about 1700 L d-1. About 10% or180 L d-1 passes into the glomerulus, or about 125 mL per minute.

    The kidney re-adsorbs all

    but 1.5 L d-1, achieving a

    100x concentration of thefiltrate.

    The corpuscles are in the

    cortex of the kidney,

    while the loops of Henle

    are lined up parallel in the

    medulla.

    KK 14.1, H&G

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    Bird KidneysBirds and mammals are the only vertebrates that can produce urine that has a salt

    concentration higher than their blood. Birds also have loops in their nephric

    tubules, but are thought to have evolved this independently.

    The organization of the bird

    kidney is very different from

    that of mammals. The loops

    are short peripherally, andlong centrally, in each

    lobule.

    KK 14.9

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    Osmoregulation and Excretion in Vertebrates 1

    Land animals always face a

    problem of water loss, and mostdrink to replace that water.

    Fish in freshwater tend to gain

    water and must conserve salt.Fish

    in salt water generally, sharksexcepted, tend to lose water also

    as they are hyposmotic to sea

    water. Some saltwater fish are

    osmoconformers and allow their

    salt content to vary so that they

    remain isosmotic.

    Relatively few fish move back

    and forth between fresh and salt

    water.

    KK 14.12

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    Osmoregulation and

    excretion in different

    vertebrates 1KK 14.10, H&G. 15.7Saltwater teleosts are hyposmotic,

    and tend to lose water. The renal

    corpuscle is small or absent, and a

    minimum of water is lost as urine.

    NH4+ or urea is lost through gills. Salt

    gained in feeding is excreted via thegills.

    Freshwater fish arehyperosmotic

    to their environment, so tend to gain

    water through their gills and in

    feeding. They actively transport saltin through their gills. The renal

    corpuscle is large, to produce copious

    dilute urine. NH4+ is lost through

    gills.

    Sharks are slightly hyperosmotic to

    seawater, which they achieve by

    accumulating urea. The renal corpuscle is

    large to eliminate water gained through gills

    and feeding. Excess salt is excreted through

    their rectal gland.

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    Osmoregulation and excretion in

    different vertebrates 2KK 14.10, H&G. 15.7

    Birds and reptiles are water conservers. They have small

    renal corpuscles, and reduce the need to produce urine by

    converting NH4+ to uric acid. Birds, like mammals, have a

    loop in the intermediate segment of the tubule.

    Marine reptiles and birds are extreme water conservers.

    When they drink water they get salt as well. They secretetheir excess salt through specialized glands of the eye,

    nasal area or mouth.

    Mammals are generally less effective at conserving water

    than reptiles and birds. They secrete NH4+ as urea, which

    is soluble, so they must produce urine. Loops of Henlehelp to minimize water loss. Marine mammals are unique

    in being able to produce urine that is hyperosmotic to sea

    water, so they can drink sea water without accumulating

    salt.

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    Summary of Pathways of Ammonia Elimination

    in Vertebrates

    KK 14.11

    Fish and aquaticreptiles

    Birds and reptiles

    Sharks,

    Sarcopterygii,

    Amphibia,

    mammals