Chapter 12 Excretory system

12.1 Evolution of Excretory Systems

Selective excretion is crucial to internal fluid homeostasis and involves

several systems

Primary functions of Excretory Systems:

1. Maintenance of proper internal levels of inorganic solutes (Na+, K+,

Cl, H+, CO2, and so forth).
2. Maintenance of proper plasma water volume.
3. Removal of nonnutritive and harmful substances resulting from
metabolism (ammonia, urea, bilirubin, and so forth) or ingestion (such
as plant alkaloids, drugs), and removal of hormones after they have
had their desired effect.
4. Maintenance of osmotic balance

Simple aquatic animals (sponges, Cnidarians, and possibly echinoderms)

- do not have specialized organs for fluid-regulating processes.
- rely on diffusion and membrane transporters for nonsolid wastes.
- Individual cells of freshwater sponges also have contractile vacuoles,
specialized vesicles that remove and eject excess water

Thus, the evolution of specialized excretory tissues was favored with

transport epithelia (linings with specific membrane mechanisms to
remove nonfecal wastes and to regulate other solutes and water).

Animal Kingdom: Four organs systems involved in excretion and retention:

1. Respiratory systems (gills and lungs), which help regulate CO2
(although we do not often think of it in these terms, the act of exhaling CO2
is a form of waste excretion). Gills can also help remove other solutes such
as ammonia and HCO3.
2. Digestive systems, which remove not only undigested food but also
some end products of internal metabolism.
3. Integument (skin) and glands, some of which can excrete organic
wastes as secondary functions (such as sweat glands in humans), and others
that evolved primarily to excrete excess inorganic ions (salt glands of brine
shrimp and marine reptiles). The skin of many amphibians also regulates salt
and water uptake.
4. Renal organs (renalis, Latin for kidney) with tubules that filter body
fluids and regulate water, ions, and many organic substances, and then
selectively reabsorb or secrete these molecules.
Nitrogen metabolism creates special stresses and produces three major end
products: ammonia, urea, and uric acid

Ammonia
- Include most bony fishes, larval amphibians, and most non
vertebrates
- rely on ammonia excretion, usually via gills. This feature is called
ammonotely.
- NH3
Urea
- usually in the liver
- transport the product to the excretory organs
- Urea excretion is termed ureotely.
- Amphibians, whose aquatic larval stages use ammonia excretion
via their gills, whereas semiaquatic adults switch to urea
production and excretion using their livers and kidneys.
- Urea is the primary nitrogenous waste.
- Urea is produced, in most vertebrates that make it, from two
ammonium ions (one via aspartic acid) and a bicarbonate ion,
using ATP, in the ornithineurea cycle, or OUC. (Some bony
fishes that produce urea use the uricolytic pathway).
- In ruminants, urea has another function. After synthesis in the
liver, it can enter the rumen via the salivary glands, where
symbiotic microbes ferment the food; there, microbes convert
urea back to ammonia, using the enzyme urease. The ammonia
can then be used directly by these beneficial microbes for their
own protein synthesis.
Uric Acid
- Uric acid is the end product of amino acid and purine (nucleic
acid) metabolism and requires even more ATP than urea to
produce.
- It is less toxic (highly insoluble) has the added benefit of
removing four nitrogens per molecule, and is excreted in a
semisolid form, generally into the hindgut.
- Serves as a protective antioxidant.
- Humans lack the enzymes needed to break down uric acid and
thus excrete uric acid as the end product of nucleic acid
metabolism.
- In any case, mammals have not evolved mechanisms to handle
high concentrations of the highly insoluble uric acid, and
excessive levels can lead to arthritic gout and kidney stones
because of crystals of this highly insoluble molecule forming in
inappropriate places. Mammals do make uric acid, but most only
synthesize small amounts that are removed by the kidneys.
Excretory organs have transport epithelia, which typically use Na+/K+
ATPases

2 transports:

Salt Transport Only

Transporting salt without water involves two steps:

- The Na+/K+ ATPase pump uses ATP to increase the Na+

concentration outside the cells basolateral side, and decrease it
inside the cell; some Na+ then moves passively into the cell from
the apical side through an ENaC channel.

- The build-up of positively charged Na+ in the basolateral fluid

passively draws Cl ions by charge attraction. Cl moves
through either or both of the following:
a. A ClC (chloride-activated chloride) channel. ClC
channels are an ancient, large family of chloride channels
found in all domains and kingdoms of life. As gated channels,
many ClC channels open in response to increasing Cl
concentration, hence their name.
b. A CFTR (cystic fibrosis transmembrane-conductance
regulator) chloride channel, so named because its mutated,
nonworking form is responsible for the defective salt transport
characteristic of cystic fibrosis. It is regulated by cAMP and
phosphorylation. Humans with cystic fibrosis have
nonfunctioning CFTRs, and cannot properly transport salt and
water through membranes of several epithelia such as linings
of airways, sweat glands, and intestine. In airways, for
example, harmful thick, sticky mucus builds up.

Overall, then, NaCl is moved across the tissue through the cells that is,
transcellularly. The tissue in is fairly waterproof, so little or no water
moves.

Salt and Water Transport

In the second tissue, the first two steps are the same, but there are
two more:
 - The osmotic pressure of transported NaCl builds up within lateral
spaces between the cells, where it induces osmosis of H2O from
the cells. The accumulation of H2O in those lateral spaces results
in a buildup of hydrostatic pressure, flushing H2O out of the
spaces into the interstitial fluid (and, in animals with blood
vessels, into nearby capillaries)
- In some cases, some water and ions are thought to move
through spaces between the cells, that is paracellularly.

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12.2 Renal Excretory Organs: Overview

Renal organs have tubules that filter solutes and water from body fluids and
subsequently modify the fluid in the tubular lumen using transport epithelia,
specifically for water and solute homeostasis and waste removal.

Renal tubules operate by two or more of four basic renal processes:

Filtration
- Solutes are selectively separated from a solution by passing through
a boundary.
- Ultrafiltration is typically driven by hydrostatic pressure, mostly a
nonselective process, where some water with all its small solutes
passes through the barrier, except that cells and large molecules
such as proteins (and some water) remain behind.
Secretion
- in which transport epithelia move (often actively) specific solutes
into the tubule lumen for excretion.
Reabsorption
- in which transport epithelia move (often actively) specific solutes
(and often water) back into the body from the lumen.
Osmoconcentration
- in which water is removed from the lumen fluid while leaving
solutes behind, producing an excretory fluid more concentrated
(hyperosmotic) than body fluids and thus conserving water.

The major renal organs:

Protonephridia (bundles of flame cells)

- use ultrafiltration driven by cilia, with secretion and reabsorption.
These are the types of specialized filtration system thought to have
 evolved first in freshwater nonvertebrates. These are generally
blind-end ducts that project into the body cavity. They have low
(fluid) hydrostatic because of the beating of cilia in the duct, which
resembles a flame, move fluid outward. The duct after the filter may
further modify the filtrate by reabsorbing desired molecules and
secreting undesired ones. The action of the cilia eventually moves
this modified fluid or urine out through a pore in the epidermis into
the external environment.
- found in more simple animals having one internal fluid
compartment, such as rotifers, flatworms, larval annelids, and larval
mollusks; they are also found in larval fishes and amphibians.

Mesonephridia and metanephridia

- use ultrafiltration typically driven by fluid pressure, with secretion,
reabsorption, and occasionally osmoconcentration.
- These tubules are in animals with two or more major fluid spaces
a coelom and a circulatory fluid.
- The tubules begin with a filtering capsule or funnel-like opening
that ultrafilter the circulatory fluid. This is followed by tubule
segments that perform selective secretion and reabsorption and
may empty via a connecting canal or ureter into a hindgut or a
bladder.
- In vertebrates, the renal tubules are called nephrons (Greek
nephros, for kidney), and they go through significant
developmental changes in most groups. In all groups, the first renal
structure is a ciliated pronephric one, which becomes functional
only in larval fishes, amphibians, adult hagfish, and lampreys. In all
other adult fishes and amphibians, the pronephric tubules
degenerate except the duct, which becomes the foundation of new
pressure driven mesonephric tubules (the kidneys of these aquatic
animals).
- Finally, in terrestrial vertebrates, both the pro and mesonephric
tubules form in embryos but are replaced by the metanephric
tubules (more adapted for water-limited life on land). The
mesonephric tubules, however, become the ducts of reproductive
organs in males. Metanephric-type tubules have evolved
independently several timesin adult mollusks, in annelids, in some
arthropods, and in vertebrates.
- In some cases, a single filtering tubule forms the renal organ, as in
the crustacean antennal gland. Each single tubule begins with a
capsule that filters hemolymph; enters a labyrinth of epithelia and
 fluid spaces followed by a coiled nephridial canal, where selective
absorption and secretion take place; and then empties into a
bladder.
- In other animals, many meso- and metanephridial tubules are
assembled together with connective tissue into large organs called
kidneys, as in the case in higher vertebrates.
Malpighian tubules
- begin filtration driven by active secretion of ions from an
arthropods hemolymph and then, in conjunction with the hindgut,
use reabsorption and sometimes osmoconcentration.

12.3 Insect Malpighian Tubules

Malpighian tubules initiate excretion by ion secretion, which causes
osmosis.
filter the hemolymph not by pressure, as in most other renal organs,
but by secreting K+ (and often Na+) into tubule lumen
The tubules and hindgut modify the lumen fluid by specific secretion
and reabsorption.

Insect excretion is regulated by diuretic and antidiuretic hormones

Diuretic hormones: In the kissing bugs and mosquitoes, which must
excrete considerable fluid loads, both serotonin and several
neuropeptides stimulate excretion.
Antidiuretic hormones: As antagonists, peptide hormones such as
cardioacceleratory peptide 2b (CAP2b) in kissing bugs and Tenmo-
ADFa in mealworm beetle larvae (Tenebrio molitor) trigger cGMP
production in principal cells, which reduces secretion and conserves
water.

12.4 Vertebrate Urinary Systems and Extrarenal Organs

Vertebrate kidneys form the urine; the remainder of the urinary system is
ductwork that carries the urine to the bladder or hindgut
The urinary system consists of the urine-forming organs
o Kidneys (2)
Nephron
functional unit of the vertebrate kidney bound
together by connective tissue.
Each nephron consists of the tubules and an
associated vascular (blood vessel) component, both
 of which are intimately related structurally and
functionally.
Consist of outer, granular-appearing renal cortex
and an inner, striated renal medulla. (small
mammals: unipapillary medulla, Larger mammals:
renal pyramids)
Each kidney is supplied by a renal artery and a renal
vein.
Vascular component
Afferent arteriolecarries blood to the
glomerulus
Glomerulusa tuft of capillaries that filters a
protein-free plasma into the tubular component
Efferent arteriolecarries blood from the
glomerulus
Peritubular capillariessupply the renal
tissue; involved in exchanges with the fluid in
the tubular lumen
Tubular component
Bowmans capsulecollects the glomerular
filtrate
Proximal tubuleuncontrolled reabsorption
and secretion of selected substances occur
here
Loop of Henleestablishes an osmotic
gradient in the renal medulla that is important
in the kidneys ability to produce urine of
varying concentration
Distal tubule and collecting duct variable,
controlled reabsorption of Na+ and H2O and
secretion of K+ and H+ occur here; fluid
leaving the collecting duct is urine, which
enters the renal pelvis
Combined vascular/tubular component
Juxtaglomerular apparatus involved in the
control of
kidney function
o Ureters (2)
These narrow tubes carry urine from the kidneys to the
bladder
Ducts walled with smooth muscle
o Bladder
o Sphincter muscles (2)
o Nerves in the bladder
o Urethra
 Present in male mammals

12.5 Mammalian Urinary System: Overview and Glomerular Filtration

Functions of mammalian kidneys:

major inorganic solutes
plasma volume
harmful or unneeded organic molecules
osmotic balance
Secretion of erythropoietin, a hormone that stimulates red blood
cell production
Secretion of renin, a hormone important for salt conservation
Conversion of vitamin D into its active form
Excretion of pheromones for sexual signaling

12.9 Mammalian Urinary System: Bladder Storage and Micturition

The mammalian bladder wall consists of smooth muscle lined by

transitional epithelium with special cells called umbrella cell.
The exit from the bladder, however, is guarded by two sphincters
o internal urethral sphincter
consists of smooth muscle and is not under cortical control
is not really a separate muscle but instead consists of the
last portion of the bladder.
Not a true sphincter
o External urethral sphincter
skeletal muscle and is under cortical control via a motor
neuron, which is normally active to keep the sphincter
closed.
Micturition (urination)
-the process of bladder emptying, is governed by two mechanisms: the
micturition spinal reflex and cortical control. The micturition reflex is initiated
when stretch receptors within the bladder wall are stimulated.