Chapter

44

Osmoregulation and
Excretion
Biology

Eighth Edition
Neil Campbell and Jane Reece

Copyright 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings

Overview: A balancing act

The physiological systems of animals
Operate in a fluid environment

The relative concentrations of water and

solutes in this environment
Must be maintained within fairly narrow limits

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Overview: A balancing act

Freshwater animals
Show adaptations that reduce water uptake and
conserve solutes

Desert and marine animals face desiccating

environments
With the potential to quickly deplete the body water

Figure 44.1
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Osmotic Challenges
Osmoconformers, which are mostly marine
invertebrates
Are isoosmotic with their surroundings and do
not regulate their osmolarity

Osmoregulators (mostly marine vertebrates)

expend energy to control water uptake and loss
In a hyperosmotic or hypoosmotic environment

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Living in a Hyperosmotic Environment -a physiological desert

Marine bony fishes are hypoosmotic to sea water

And lose water by osmosis and gain salt by both
diffusion and from food they eat

These fishes balance water loss

By drinking seawater
Gain of water and
salt ions from food
and by drinking
seawater

Excretion of
salt ions
from gills

Figure 44.3a
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Osmotic water loss

through gills and other parts
of body surface

Excretion of salt ions

and small amounts
of water in scanty
urine from kidneys

(a) Osmoregulation in a saltwater fish

Living in a Hypoosmotic Environment

Freshwater animals maintain water balance
By excreting large amounts of dilute urine

Salts lost by diffusion

Are replaced by foods and uptake across the gills
Uptake of
water and some
ions in food

Osmotic water gain

through gills and other parts
of body surface

Uptake of
salt ions
by gills

Excretion of
large amounts of
water in dilute
urine from kidneys

Figure 44.3b (b) Osmoregulation in a freshwater fish

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Land Animals
Land animals
manage their
water budgets

Water
balance in
a human
(2,500 mL/day
= 100%)

Water
balance in a
kangaroo rat
(2 mL/day
= 100%)

By drinking and
eating moist
foods and by
using metabolic
water

Ingested
in food (750)

Ingested
in food (0.2)
Ingested
in liquid
(1,500)

Water
gain

Derived from
metabolism (250)

Derived from
metabolism (1.8)

Feces (0.9)
Urine
(0.45)

Urine
(1,500)

Water
loss

Figure 44.5
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Feces (100)

Evaporation (1.46)

Evaporation (900)

Kangaroo Rat (Dipodomys ordii)

From Animal Physiology, by Roger Eckert, David Randall, and George Augustine, Copyright 1998 by W. H. Freeman
and Company. Used with permission.

Animal
Freshwater
fish

Inflow/Outflow
Does not drink
water H2O in
Salt in
(active transport by gills)

Urine
Large volume
of urine
Urine is less
concentrated
than body
fluids

Salt out
Bony marine
fish

Drinks water
Salt in H2O out

Small volume
of urine
Urine is
slightly less
concentrated
than body
fluids

Salt out (active

transport by gills)
Terrestrial
vertebrate

Drinks water
Salt in
(by mouth)

H2O and
salt out

Moderate
volume
of urine
Urine is
more
concentrated
than body
fluids

Nitrogenous Wastes
Among the most
important wastes
Are the
nitrogenous
breakdown
products of
proteins and
nucleic acids

Nucleic acids

Proteins

Nitrogenous bases

Amino acids

NH2
Amino groups

Most aquatic
animals, including
most bony fishes

Many reptiles
(including
birds), insects,
land snails

Mammals, most
amphibians, sharks,
some bony fishes

O
C
NH2
NH3
Ammonia

C
NH2

HN

Urea

Figure 44.9
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H
N
C

N
H

N
H

Uric acid

Ammonia
Animals that excrete
nitrogenous wastes as
ammonia

Nitrogenous bases

Amino acids

NH2
Amino groups

Need access to lots of

water
Release it across the
whole body surface or
through the gills

Nucleic acids

Proteins

Most aquatic
animals, including
most bony fishes

Many reptiles
(including
birds), insects,
land snails

Mammals, most
amphibians, sharks,
some bony fishes
O
C
NH2

NH3
Ammonia

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

C
NH2
Urea

HN

N
H

N
C
N
H

Uric acid

Urea
The liver of mammals
and most adult
amphibians

Nitrogenous bases

Amino acids

NH2
Amino groups

Converts ammonia to
less toxic urea

Urea is carried to the

kidneys, concentrated

Nucleic acids

Proteins

Most aquatic
animals, including
most bony fishes

Many reptiles
(including
birds), insects,
land snails

Mammals, most
amphibians, sharks,
some bony fishes
O

And excreted with a

minimal loss of water

NH3
Ammonia

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

NH2
NH2
Urea

HN

N
H

H
N
C
N
H

Uric acid

Uric Acid
Insects, land snails, and
many reptiles, including
birds

Nitrogenous bases

Amino acids

NH2
Amino groups

Excrete uric acid as their

major nitrogenous waste

Uric acid is largely

insoluble in water

Nucleic acids

Proteins

Most aquatic
animals, including
most bony fishes

Many reptiles
(including
birds), insects,
land snails

Mammals, most
amphibians, sharks,
some bony fishes
O

And can be secreted as

a paste with little water
loss
Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

NH3
Ammonia

NH2
NH2
Urea

HN

N
H

H
N
C
N
H

Uric acid

Excretory Processes
Most excretory systems
Produce urine by refining a filtrate derived from
body fluids
Capillary

Filtrate

Excretory
tubule

1 Filtration. The excretory tubule collects a filtrate from the blood.

Water and solutes are forced by blood pressure across the
selectively permeable membranes of a cluster of capillaries and
into the excretory tubule.

2 Reabsorption. The transport epithelium reclaims valuable substances

from the filtrate and returns them to the body fluids.

3 Secretion. Other substances, such as toxins and excess ions, are

extracted from body fluids and added to the contents of the excretory
tubule.

Urine

Figure 44.10
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4 Excretion. The filtrate leaves the system and the body.

Protonephridia: Flame-Bulb Systems

A protonephridium

Is a network of deadend tubules lacking

internal openings

Nucleus
of cap cell

The tubules branch

throughout the body

Interstitial fluid
filters through
membrane where
cap cell and tubule
cell interdigitate
(interlock)

And the smallest

branches are capped
by a cellular unit called
a flame bulb

These tubules excrete a

dilute fluid

Cilia

And function in
osmoregulation

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Tubule cell

Flame
bulb
Protonephridia
(tubules)

Figure 44.11

Tubule

Nephridiopore
in body wall

Metanephridia
Each segment of an earthworm
Has a pair of open-ended metanephridia

Coelom
Capillary
network
Bladder
Collecting
tubule

Nephridiopore

Figure 44.12

Nephrostome

Metanephridia

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Malpighian Tubules
In insects and other terrestrial arthropods,
malpighian tubules
Remove nitrogenous wastes from hemolymph
and function in osmoregulation
Digestive tract

Rectum
Intestine

Hindgut

Malpighian
Midgut
tubules
(stomach)
Salt, water, and
nitrogenous
wastes

Feces and urine

Anus

Malpighian
tubule
Rectum

Figure 44.13

HEMOLYMPH

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Reabsorption of H2O,
ions, and valuable
organic molecules

Vertebrate Kidneys
Kidneys, the excretory
organs of vertebrates
function in both
excretion and
osmoregulation and
are often paired

Each kidney is supplied

with blood by a renal
artery and drained by a
renal vein
Nephrons and associated
blood vessels are the
functional unit of the
mammalian kidney
Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

Posterior vena cava

Renal artery and vein

Kidney

Aorta
Ureter
Urinary bladder
Urethra
(a) Excretory organs and major
associated blood vessels

Structure and Function of the Nephron and Associated

Structures

The mammalian kidney has two distinct regions

An outer renal cortex and an inner renal
medulla
Renal
medulla
Renal
cortex
Renal
pelvis

Ureter

Figure 44.14b

Section of kidney from a rat

(b) Kidney structure

Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

The Nephron
The nephron, the functional unit of the vertebrate
kidney
Consists of a single long tubule and a ball of
capillaries called the glomerulus
JuxtaCortical
medullary nephron
nephron

Afferent
arteriole
from renal
artery

Glomerulus
Bowmans capsule
Proximal tubule

Renal
cortex

Peritubular
capillaries
Collecting
duct

To
renal
pelvis

20 m
Renal
medulla

SEM
Efferent
arteriole from
glomerulus

Loop
of
Henle

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Collecting
duct

Branch of
renal vein
Descending
limb
Ascending
limb

Figure 44.13c, d
(c) Nephron

Distal
tubule

(d) Filtrate and

blood flow

Vasa
recta

From Blood Filtrate to Urine: A Closer Look

Filtrate becomes urine
As it flows through the mammalian nephron
and collecting duct
1 Proximal tubule
NaCl Nutrients
HCO3
H2 O
K+

H+

NH3

4 Distal tubule
NaCl

H2 O

K+

HCO3

H+

CORTEX
Filtrate
H2 O
Salts (NaCl and others)
HCO3
H+
Urea
Glucose; amino acids
Some drugs

2 Descending limb
of loop of
Henle

3 Thick segment
of ascending
limb
NaCl

H2O
OUTER
MEDULLA

NaCl
3 Thin segment
of ascending
limb

Key
Active transport
Passive transport

5 Collecting
duct
Urea

NaCl
INNER
MEDULLA

Figure 44.15
Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings

H2O