Excretory System

KK Chapter 14, Hildebrand Chapter 15

Excretory System Functions

1) Maintain salt/water balance by releasing controlled amounts of both under the influence of the endocrine system (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 in most fish as NH4,+ but otherwise the liver may turn NH4+ into less toxic urea. In birds and reptiles, the kidneys turn NH4+ into uric acid.

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 passes from the blood-vascular system to the capsule, while the tubule reclaims solutes from the filtrate and conducts urine towards the outside of the body.

Development of the Kidney

KK 14.3a, H&G 15.1 The kidney tissue develops from the the 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 the mesomere 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 a glomerulus, and each is associated with a capsule and tubule.

Development of the Kidney 2

KK 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. In amniotes, 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

Development of the Kidney 3

KK 14.4, H&G 15.2

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 in non-amniotes, and is then called the opisthonephros (tail kidney). In amniotes, a third kidney or metanephros develops behind the embryonic 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.
KK 14.4

Amniote embryo illustrating all 3 kidney types

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 we shall see, thats not all!

KK 14.5, H&G 15.2

Opisthonephric Kidney of a (Male) Salamander

There are many renal corpuscles 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 KK 14.8

Mammalian Kidney Function

The blood supply to the glomerulus is from the renal artery. 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 actively removed, and Cl- water follows passively.
KK 14.14, H&G 15.8

Mammalian Kidney Function 2

The intermediate part of the tubule is a site of active tranport of Na+ into the surronding tissue, resulting in a very high Na+ concentration in that tissue. Clfollows 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.

Human Metanephric Kidney

The 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% or 180 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 the filtrate. 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

Bird Kidneys
Birds 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, and long centrally, in each lobule.

KK 14.9

Osmoregulation and Excretion in Vertebrates 1

Land animals always face a problem of water loss, and most drink to replace that water. Fish in freshwater tend to gain water and must conserve salt. Fish in salt water generally, sharks excepted, 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

Osmoregulation and excretion in different vertebrates 1 KK 14.10, H&G. 15.7

Saltwater 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 the gills. Freshwater fish are hyperosmotic to their environment, so tend to gain water through their gills and in feeding. They actively transport salt in 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.

Osmoregulation and excretion in different vertebrates 2 KK 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 secrete their 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 Henle help 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.

Summary of Pathways of Ammonia Elimination in Vertebrates

Birds and reptiles

Sharks, Sarcopterygii, Amphibia, mammals Fish and aquatic reptiles

KK 14.11