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Chapter 44

Osmoregulation and
Excretion

PowerPoint Lectures for


Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero


Copyright © 2005 Pearson Education, Inc. publishing as 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

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• 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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• Osmoregulation
– Regulates solute concentrations and balances
the gain and loss of water

• Excretion
– Gets rid of metabolic wastes

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• Concept 44.1: Osmoregulation balances the
uptake and loss of water and solutes
• Osmoregulation is based largely on controlled
movement of solutes
– Between internal fluids and the external
environment

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Osmosis
• Cells require a balance
– Between osmotic gain and loss of water

• Water uptake and loss


– Are balanced by various mechanisms of
osmoregulation in different environments

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Osmotic Challenges
• Osmoconformers, which are only marine
animals
– Are isoosmotic with their surroundings and do
not regulate their osmolarity

• Osmoregulators expend energy to control


water uptake and loss
– In a hyperosmotic or hypoosmotic environment

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• Most animals are said to be stenohaline
– And cannot tolerate substantial changes in
external osmolarity

• Euryhaline animals
– Can survive large fluctuations in external
osmolarity

Figure 44.2
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Marine Animals
• Most marine invertebrates are osmoconformers
• Most marine vertebrates and some
invertebrates are osmoregulators

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• 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
Osmotic water loss
salt ions from food
through gills and other parts
and by drinking
of body surface
seawater

Excretion of Excretion of salt ions


salt ions and small amounts
from gills of water in scanty
urine from kidneys

Figure 44.3a (a) Osmoregulation in a saltwater fish


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Freshwater Animals
• Freshwater animals
– Constantly take in water from their
hypoosmotic environment
– Lose salts by diffusion

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• 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
Osmotic water gain
through gills and other parts
of body surface
Uptake of
water and some
ions in food

Uptake of Excretion of
salt ions large amounts of
by gills water in dilute
urine from kidneys

Figure 44.3b (b) Osmoregulation in a freshwater fish


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Animals That Live in Temporary Waters
• Some aquatic invertebrates living in temporary
ponds
– Can lose almost all their body water and
survive in a dormant state

• This adaptation is called anhydrobiosis 100 µm

100 µm

Figure 44.4a, b (a) Hydrated tardigrade (b) Dehydrated tardigrade

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Land Animals
• Land animals manage their water budgets
– By drinking and eating moist foods and by
using metabolic water Water
balance in a
kangaroo rat
Water
balance in
a human
(2 mL/day (2,500 mL/day
= 100%) = 100%)

Ingested Ingested
in food (0.2) in food (750)

Ingested
Water in liquid
gain (1,500)

Derived from Derived from


metabolism (1.8) metabolism (250)

Feces (0.9) Feces (100)


Urine Urine
(0.45) (1,500)
Water
loss

Evaporation (1.46) Evaporation (900)


Figure 44.5
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• Desert animals
– Get major water savings from simple anatomical
features
EXPERIMENT Knut and Bodil Schmidt-Nielsen and their colleagues from Duke University observed that the
fur of camels exposed to full sun in the Sahara Desert could reach temperatures of over 70°C, while the
animals’ skin remained more than 30°C cooler. The Schmidt-Nielsens reasoned that insulation of the skin
by fur may substantially reduce the need for evaporative cooling by sweating. To test this hypothesis, they
compared the water loss rates of unclipped and clipped camels.

RESULTS Removing the fur of a camel increased the rate


of water loss through sweating by up to 50%.
4

(L/100 kg body mass)


Water lost per day
3

0
CONCLUSION The fur of camels plays a critical role in
Control group Experimental group
their conserving water in the hot desert (Unclipped fur) (Clipped fur)
environments where they live.

Figure 44.6
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Transport Epithelia
• Transport epithelia
– Are specialized cells that regulate solute
movement
– Are essential components of osmotic
regulation and metabolic waste disposal
– Are arranged into complex tubular networks

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• An example of transport epithelia is found in the
salt glands of marine birds
– Which remove excess sodium chloride from the
Nasal salt gland
blood
(a) An albatross’s salt glands Nostril
empty via a duct into the with salt
nostrils, and the salty solution secretions
either drips off the tip of the
beak or is exhaled in a fine mist.
Lumen of
secretory tubule
Vein
Capillary
(c) The secretory cells actively
Secretory
transport salt from the
tubule Artery
blood into the tubules.
NaCl
Transport Blood flows counter to the
epithelium flow of salt secretion. By
(b) One of several thousand
maintaining a concentration
secretory tubules in a salt-
Direction gradient of salt in the tubule
excreting gland. Each tubule
of salt (aqua), this countercurrent
is lined by a transport
movement Blood Secretory cell system enhances salt
epithelium surrounded by
flow transfer from the blood to
capillaries, and drains into of transport
the lumen of the tubule.
a central duct. epithelium
Central
Figure 44.7a, b duct

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• Concept 44.2: An animal’s nitrogenous wastes
reflect its phylogeny and habitat
• The type and quantity of an animal’s waste
products
– May have a large impact on its water balance

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• Among the most important wastes
– Are the nitrogenous breakdown products of
proteins and nucleic acids
Proteins Nucleic acids

Amino acids Nitrogenous bases

–NH2
Amino groups

Many reptiles
Most aquatic Mammals, most (including
animals, including amphibians, sharks,
birds), insects,
most bony fishes some bony fishes
land snails

O
H
C
HN C N
NH2 C O
O C C C N
NH3 O N
NH2 H H
Ammonia Urea Uric acid
Figure 44.8
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Forms of Nitrogenous Wastes
• Different animals
– Excrete nitrogenous wastes in different forms

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Ammonia
• Animals that excrete nitrogenous wastes as
ammonia
– Need access to lots of water
– Release it across the whole body surface or
through the gills

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Urea
• The liver of mammals and most adult
amphibians
– Converts ammonia to less toxic urea

• Urea is carried to the kidneys, concentrated


– And excreted with a minimal loss of water

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Uric Acid
• Insects, land snails, and many reptiles,
including birds
– Excrete uric acid as their major nitrogenous
waste

• Uric acid is largely insoluble in water


– And can be secreted as a paste with little
water loss

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The Influence of Evolution and Environment on
Nitrogenous Wastes

• The kinds of nitrogenous wastes excreted


– Depend on an animal’s evolutionary history
and habitat

• The amount of nitrogenous waste produced


– Is coupled to the animal’s energy budget

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• Concept 44.3: Diverse excretory systems are
variations on a tubular theme
• Excretory systems
– Regulate solute movement between internal
fluids and the external environment

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Excretory Processes
• Most excretory systems
– Produce urine by refining a filtrate derived from
body fluids
Capillary
1 Filtration. The excretory tubule collects a filtrate from the blood.
Water and solutes are forced by blood pressure across the
Excretory
selectively permeable membranes of a cluster of capillaries and

Filtrate
tubule
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

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

Figure 44.9
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• Key functions of most excretory systems are
– Filtration, pressure-filtering of body fluids
producing a filtrate
– Reabsorption, reclaiming valuable solutes from
the filtrate
– Secretion, addition of toxins and other solutes
from the body fluids to the filtrate
– Excretion, the filtrate leaves the system

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Survey of Excretory Systems
• The systems that perform basic excretory
functions
– Vary widely among animal groups
– Are generally built on a complex network of
tubules

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Protonephridia: Flame-Bulb Systems
• A protonephridium
– Is a network of dead-end tubules lacking
internal openings Nucleus
of cap cell

Cilia

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

Flame
bulb

Protonephridia
(tubules) Tubule

Nephridiopore
Figure 44.10 in body wall

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• The tubules branch throughout the body
– And the smallest branches are capped by a
cellular unit called a flame bulb

• These tubules excrete a dilute fluid


– And function in osmoregulation

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

Coelom
Capillary
network

Bladder
Collecting
tubule

Nephridio-
pore
Figure 44.11 Nephrostome Metanephridia
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• Metanephridia consist of tubules
– That collect coelomic fluid and produce dilute
urine for excretion

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

Rectum
Hindgut
Intestine

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

Malpighian
tubule
Rectum

Reabsorption of H2O,
ions, and valuable
organic molecules
HEMOLYMPH
Figure 44.12
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• Insects produce a relatively dry waste matter
– An important adaptation to terrestrial life

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Vertebrate Kidneys
• Kidneys, the excretory organs of vertebrates
– Function in both excretion and osmoregulation

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• Concept 44.4: Nephrons and associated blood
vessels are the functional unit of the
mammalian kidney
• The mammalian excretory system centers on
paired kidneys
– Which are also the principal site of water
balance and salt regulation

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• Each kidney
– Is supplied with blood by a renal artery and
drained by a renal vein

Posterior vena cava

Renal artery and vein

Aorta Kidney

Ureter

Urinary bladder

Urethra

(a) Excretory organs and major


associated blood vessels
Figure 44.13a
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• Urine exits each kidney
– Through a duct called the ureter

• Both ureters
– Drain into a common urinary bladder

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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
Section of kidney from a rat
Figure 44.13b (b) Kidney structure

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• The nephron, the functional unit of the vertebrate
kidney
– Consists of a single long tubule and a ball of
capillaries called the glomerulus
Juxta- Cortical
medullary nephron
nephron Afferent
arteriole
from renal Glomerulus
artery
Bowman’s capsule
Renal
cortex Proximal tubule
Peritubular
capillaries

Collecting
SEM
duct 20 µm
Efferent Distal
Renal
arteriole from tubule
To medulla glomerulus
renal
pelvis Branch of Collecting
renal vein duct
Descending
Loop limb
of
Ascending
Henle
limb

Figure 44.13c, d Vasa


(d) Filtrate and recta
(c) Nephron
blood flow
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Filtration of the Blood
• Filtration occurs as blood pressure
– Forces fluid from the blood in the glomerulus
into the lumen of Bowman’s capsule

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• Filtration of small molecules is nonselective
– And the filtrate in Bowman’s capsule is a
mixture that mirrors the concentration of
various solutes in the blood plasma

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Pathway of the Filtrate
• From Bowman’s capsule, the filtrate passes
through three regions of the nephron
– The proximal tubule, the loop of Henle, and the
distal tubule

• Fluid from several nephrons


– Flows into a collecting duct

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Blood Vessels Associated with the Nephrons
• Each nephron is supplied with blood by an afferent
arteriole
– A branch of the renal artery that subdivides into the
capillaries

• The capillaries converge as they leave the glomerulus


– Forming an efferent arteriole

• The vessels subdivide again


– Forming the peritubular capillaries, which surround the
proximal and distal tubules

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From Blood Filtrate to Urine: A Closer Look
• Filtrate becomes urine
– As it flows through the mammalian nephron
and collecting duct
1 Proximal tubule 4 Distal tubule
NaCl Nutrients H2O
HCO3 H2O K+ NaCl HCO3

H+ NH3 K+ H+

CORTEX
2 Descending limb 3 Thick segment
Filtrate of loop of of ascending
H2O Henle limb
Salts (NaCl and others)
NaCl
HCO3–
H2O
H+
Urea OUTER NaCl
Glucose; amino acids MEDULLA
Some drugs 3 Thin segment 5 Collecting
of ascending duct
limb
Key Urea

Active transport NaCl H2O


Passive transport INNER
MEDULLA

Figure 44.14
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• Secretion and reabsorption in the proximal
tubule
– Substantially alter the volume and composition
of filtrate

• Reabsorption of water continues


– As the filtrate moves into the descending limb
of the loop of Henle

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• As filtrate travels through the ascending limb of the
loop of Henle
– Salt diffuses out of the permeable tubule into the
interstitial fluid
• The distal tubule
– Plays a key role in regulating the K+ and NaCl
concentration of body fluids
• The collecting duct
– Carries the filtrate through the medulla to the renal
pelvis and reabsorbs NaCl

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• Concept 44.5: The mammalian kidney’s ability
to conserve water is a key terrestrial adaptation
• The mammalian kidney
– Can produce urine much more concentrated
than body fluids, thus conserving water

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Solute Gradients and Water Conservation
• In a mammalian kidney, the cooperative action
and precise arrangement of the loops of Henle
and the collecting ducts
– Are largely responsible for the osmotic
gradient that concentrates the urine

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• Two solutes, NaCl and urea, contribute to the
osmolarity of the interstitial fluid
– Which causes the reabsorption of water in the
kidney and concentrates the urine
Osmolarity of
interstitial
fluid
300 (mosm/L)
300 100
300
100
300 300
CORTEX H2O NaCl H2O
Active
transport 400 200 400 400
H2O NaCl H2O
Passive
transport
H2O NaCl H2O

OUTER H2O NaCl H2O


MEDULLA 600 400 600 600
H2O NaCl H2O
Urea
H2O NaCl H2O 900
900 700
Urea
INNER H2O NaCl H2O
MEDULLA Urea 1200
1200
1200

Figure 44.15
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• The countercurrent multiplier system involving
the loop of Henle
– Maintains a high salt concentration in the
interior of the kidney, which enables the kidney
to form concentrated urine

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• The collecting duct, permeable to water but not
salt
– Conducts the filtrate through the kidney’s
osmolarity gradient, and more water exits the
filtrate by osmosis

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• Urea diffuses out of the collecting duct
– As it traverses the inner medulla

• Urea and NaCl


– Form the osmotic gradient that enables the
kidney to produce urine that is hyperosmotic to
the blood

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Regulation of Kidney Function
• The osmolarity of the urine
– Is regulated by nervous and hormonal control
of water and salt reabsorption in the kidneys

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• Antidiuretic hormone (ADH)
– Increases water reabsorption in the distal
tubules and collecting ducts of the kidney
Osmoreceptors
in hypothalamus Thirst
Hypothalamus

Drinking reduces
blood osmolarity
to set point
ADH
Increased
permeability

Pituitary
gland

Distal
tubule

H2O reab-
sorption helps
prevent further
STIMULUS:
osmolarity
The release of ADH is
increase
triggered when osmo-
receptor cells in the Collecting duct
hypothalamus detect an
increase in the osmolarity
of the blood

Homeostasis:
Blood osmolarity

(a) Antidiuretic hormone (ADH) enhances fluid retention by making


Figure 44.16a the kidneys reclaim more water.

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• The renin-angiotensin-aldosterone system (RAAS)
– Is part of a complex feedback circuit that functions
in homeostasis
Homeostasis:
Blood pressure,
volume
Increased Na+
and H2O reab-
sorption in
distal tubules

STIMULUS:
The juxtaglomerular
apparatus (JGA) responds
to low blood volume or
Aldosterone blood pressure (such as due
to dehydration or loss of
blood)

Arteriole
constriction
Adrenal gland

Angiotensin II
Distal
tubule

Angiotensinogen

JGA

Renin
production

Renin

(b) The renin-angiotensin-aldosterone system (RAAS) leads to an increase


Figure 44.16b in blood volume and pressure.

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• Another hormone, atrial natriuretic factor (ANF)
– Opposes the RAAS

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• The South American vampire bat, which feeds
on blood
– Has a unique excretory system in which its
kidneys offload much of the water absorbed from
a meal by excreting large amounts of dilute urine

Figure 44.17
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• Concept 44.6: Diverse adaptations of the
vertebrate kidney have evolved in different
environments
• The form and function of nephrons in various
vertebrate classes
– Are related primarily to the requirements for
osmoregulation in the animal’s habitat

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• Exploring environmental adaptations of the
vertebrate kidney
MAMMALS BIRDS AND OTHER REPTILES

Bannertail Kangaroo rat


(Dipodomys spectabilis) Roadrunner
(Geococcyx californianus)

Desert iguana
Beaver (Castor canadensis)
(Dipsosaurus dorsalis)
FRESHWATER FISHES AND AMPHIBIANS
MARINE BONY FISHES

Rainbow trout
(Oncorrhynchus mykiss) Northern bluefin tuna (Thunnus thynnus)

Figure 44.18 Frog (Rana temporaria)

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