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Copyright 2005 Pearson Education, Inc. publishing as Benjamin Cummings
PowerPoint Lectures forBio logy, Seventh Edit io n
Neil Campbell and Jane Reece
Lectures by Chris Romero
Chapter 44
Osmoregulation and
Excretion
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Overview: A balancing act
The physiological systems of animals
Operate in a fluid environment
The relative concentrations of water andsolutes in this environment
Must be maintained within fairly narrow limits
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Freshwater animals
Show adaptations that reduce water uptake andconserve 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 balancesthe 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 ofosmoregulation 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 controlwater 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 inexternal 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 someinvertebrates are osmoregulators
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Marine bony fishes are hypoosmotic to sea water
And lose water by osmosis and gain salt by bothdiffusion and from food they eat
These fishes balance water loss
By drinking seawater
Figure 44.3a
Gain of water and
salt ions from food
and by drinking
seawater
Osmotic water loss
through gills and other parts
of body surface
Excretion of
salt ions
from gills
Excretion of salt ions
and small amounts
of water in scanty
urine from kidneys
(a) Osmoregulation in a saltwater fish
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Freshwater Animals Freshwater animals
Constantly take in water from theirhypoosmotic 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
Figure 44.3b
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
(b) Osmoregulation in a freshwater fish
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Animals That L ive 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
Figure 44.4a, b (a) Hydrated tardigrade (b) Dehydrated tardigrade
100 m
100 m
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Land Animals Land animals manage their water budgets
By drinking and eating moist foods and byusing metabolic water
Figure 44.5
Water
balance in
a human
(2,500 mL/day
= 100%)
Water
balance in a
kangaroo rat
(2 mL/day
= 100%)
Ingested
in food (0.2)
Ingested
in food (750)
Ingested
in liquid
(1,500)
Derived from
metabolism (250)Derived from
metabolism (1.8)
Water
gain
Feces (0.9)
Urine
(0.45)
Evaporation (1.46)
Feces (100)
Urine
(1,500)
Evaporation (900)
Water
loss
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Desert animals
Get major water savings from simple anatomicalfeatures
Figure 44.6
Control group
(Unclipped fur)Experimental group
(Clipped fur)
4
3
2
1
0Wat
erlostperday
(L/100kgbodymass)
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 70C, while the
animals skin remained more than 30C 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.
EXPERIMENT
RESULTS Removing the fur of a camel increased the rate
of water loss through sweating by up to 50%.
The fur of camels plays a critical role in
their conserving water in the hot desert
environments where they live.
CONCLUSION
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Transport Epithelia
Transport epithelia
Are specialized cells that regulate solutemovement
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 theblood
Figure 44.7a, b
Nasal salt gland
Nostril
with salt
secretions
Lumen of
secretory tubule
NaCl
Blood
flowSecretory cell
of transport
epithelium
Central
duct
Direction
of salt
movement
Transportepithelium
Secretory
tubule
Capillary
Vein
Artery
(a)An albatrosss salt glands
empty via a duct into the
nostrils, and the salty solution
either drips off the tip of the
beak or is exhaled in a fine mist.
(b) One of several thousand
secretory tubules in a salt-
excreting gland. Each tubule
is lined by a transport
epithelium surrounded by
capillaries, and drains into
a central duct.
(c) The secretory cells actively
transport salt from the
blood into the tubules.
Blood flows counter to theflow of salt secretion. By
maintaining a concentration
gradient of salt in the tubule
(aqua), this countercurrent
system enhances salt
transfer from the blood to
the lumen of the tubule.
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Concept 44.2: An animals nitrogenous wastes
reflect its phylogeny and habitat The type and quantity of an animals waste
products
May have a large impact on its water balance
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Proteins Nucleic acids
Amino acids Nitrogenous bases
NH2
Amino groups
Most aquaticanimals, including
most bony fishes
Mammals, mostamphibians, sharks,
some bony fishes
Many reptiles
(includingbirds), insects,
land snails
Ammonia Urea Uric acid
NH3 NH2
NH2O C
C
CN
CO N
H H
C O
NC
HN
OH
Among the most important wastes
Are the nitrogenous breakdown products ofproteins and nucleic acids
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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Ur ic 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 onNitrogenous Wastes
The kinds of nitrogenous wastes excreted Depend on an animals evolutionary history
and habitat
The amount of nitrogenous waste produced
Is coupled to the animals 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
E P
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Excretory Processes
Most excretory systems
Produce urine by refining a filtrate derived frombody fluids
Figure 44.9
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.
Reabsorption.The transport epithelium reclaims valuable substances
from the filtrate and returns them to the body fluids.
Secretion.Other substances, such as toxins and excess ions, are
extracted from body fluids and added to the contents of the excretorytubule.
Excretion.The filtrate leaves the system and the body.
Capillary
Excretory
tubule
Filt
rate
Urine
1
2
3
4
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Key functions of most excretory systems are
Filtration, pressure-filtering of body fluidsproducing 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
S f E t S t
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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
P t h idi F l B lb S t
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Nucleus
of cap cell
Cilia
Interstitial fluid
filters through
membrane where
cap cell and tubule
cell interdigitate
(interlock)
Tubule cell
Flame
bulb
Nephridiopore
in body wall
Tubule
Protonephridia
(tubules)
Protonephr idia: F lame-Bulb Systems A protonephridium
Is a network of dead-end tubules lackinginternal openings
Figure 44.10
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The tubules branch throughout the body
And the smallest branches are capped by acellular unit called a flame bulb
These tubules excrete a dilute fluid
And function in osmoregulation
M t h idi
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Metanephr idia Each segment of an earthworm
Has a pair of open-ended metanephridia
Figure 44.11 Nephrostome Metanephridia
Nephridio-
pore
Collecting
tubule
Bladder
Capillary
network
Coelom
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Metanephridia consist of tubules
That collect coelomic fluid and produce diluteurine for excretion
M l i hi T b l
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Digestive tract
Midgut
(stomach)
Malpighian
tubules
Rectum
IntestineHindgut
Salt, water, and
nitrogenous
wastes
Feces and urineAnus
Malpighian
tubule
Rectum
Reabsorption of H2O,
ions, and valuable
organic moleculesHEMOLYMPH
Malpighian Tubules In insects and other terrestrial arthropods,
malpighian tubules
Remove nitrogenous wastes from hemolymph
and function in osmoregulation
Figure 44.12
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Insects produce a relatively dry waste matter
An important adaptation to terrestrial life
Vertebrate Kidneys
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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 anddrained by a renal vein
Figure 44.13a
Posterior vena cava
Renal artery and vein
Aorta
Ureter
Urinary bladder
Urethra
(a) Excretory organs and major
associated blood vessels
Kidney
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Urine exits each kidney
Through a duct called the ureter
Both ureters
Drain into a common urinary bladder
Structure and Function of the Nephron and Associated
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(b) Kidney structure
UreterSection of kidney from a rat
Renal
medulla
Renal
cortex
Renal
pelvis
Figure 44.13b
Structure and Function of the Nephron and AssociatedStructures
The mammalian kidney has two distinct regions An outer renal cortex and an inner renal
medulla
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The nephron, the functional unit of the vertebrate
kidney
Consists of a single long tubule and a ball ofcapillaries called the glomerulus
Figure 44.13c, d
Juxta-
medullary
nephron
Cortical
nephron
Collecting
duct
Torenal
pelvis
Renal
cortex
Renal
medulla
20 m
Afferent
arteriole
from renalartery
Glomerulus
Bowmans capsule
Proximal tubule
Peritubular
capillaries
SEM
Efferent
arteriole from
glomerulus
Branch of
renal vein
Descending
limb
Ascending
limb
Loop
of
Henle
Distal
tubule
Collecting
duct
(c) Nephron
Vasa
recta(d) Filtrate and
blood flow
F iltration of the Blood
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F iltration of the Blood Filtration occurs as blood pressure
Forces fluid from the blood in the glomerulusinto the lumen of Bowmans capsule
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Filtration of small molecules is nonselective
And the filtrate in Bowmans capsule is amixture that mirrors the concentration of
various solutes in the blood plasma
Pathway of the F iltrate
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Pathway of the F iltrate From Bowmans 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
Blood Vessels Associated with the Nephrons
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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 theproximal and distal tubules
From Blood Filtrate to Urine: A Closer Look
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Proximal tubule
Filtrate
H2O
Salts (NaCl and others)
HCO3
H+
Urea
Glucose; amino acids
Some drugs
Key
Active transport
Passive transport
CORTEX
OUTER
MEDULLA
INNER
MEDULLA
Descending limb
of loop of
Henle
Thick segment
of ascending
limb
Thin segmentof ascending
limb
Collecting
duct
NaCl
NaCl
NaCl
Distal tubule
NaCl Nutrients
Urea
H2O
NaCl
H2OH2OHCO3
K+
H+ NH3
HCO3
K+ H+
H2O
1 4
32
3 5
From Blood Filtrate to Urine: A Closer Look
Filtrate becomes urine
As it flows through the mammalian nephronand collecting duct
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 theloop of Henle
Salt diffuses out of the permeable tubule into theinterstitial fluid
The distal tubule
Plays a key role in regulating the K+and NaClconcentration of body fluids
The collecting duct
Carries the filtrate through the medulla to the renalpelvis and reabsorbs NaCl
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Concept 44.5: The mammalian kidneys ability
to conserve water is a key terrestrial adaptation
The mammalian kidney
Can produce urine much more concentrated
than body fluids, thus conserving water
Solute Gradients and Water Conservation
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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
Figure 44.15
H2O
H2O
H2O
H2O
H2O
H2O
H2O
NaCl
NaCl
NaCl
NaCl
NaCl
NaCl
NaCl
300
300 100
400
600
900
1200
700
400
200
100
Active
transport
Passive
transport
OUTERMEDULLA
INNER
MEDULLA
CORTEX
H2O
Urea
H2O
Urea
H2O
Urea
H2O
H2O
H2O
H2O
1200
1200
900
600
400
300
600
400
300
Osmolarity of
interstitial
fluid(mosm/L)300
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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 kidneys
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 thekidney to produce urine that is hyperosmotic to
the blood
Regulation of Kidney Function
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egu at o o d ey u ct o
The osmolarity of the urine
Is regulated by nervous and hormonal controlof water and salt reabsorption in the kidneys
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Antidiuretic hormone (ADH)
Increases water reabsorption in the distaltubules and collecting ducts of the kidney
Figure 44.16a
Osmoreceptors
in hypothalamus
Drinking reduces
blood osmolarity
to set point
H2O reab-
sorption helps
prevent further
osmolarity
increase
STIMULUS:
The release of ADH istriggered when osmo-
receptor cells in the
hypothalamus detect an
increase in the osmolarity
of the blood
Homeostasis:
Blood osmolarity
Hypothalamus
ADH
Pituitary
gland
Increased
permeability
Thirst
Collecting duct
Distal
tubule
(a)Antidiuretic hormone (ADH) enhances fluid retention by making
the kidneys reclaim more water.
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The renin-angiotensin-aldosterone system (RAAS)
Is part of a complex feedback circuit that functionsin homeostasis
Figure 44.16b
Increased Na+
and H2O reab-
sorption in
distal tubules
Homeostasis:
Blood pressure,
volume
STIMULUS:
The juxtaglomerular
apparatus (JGA) respondsto low blood volume or
blood pressure (such as due
to dehydration or loss of
blood)
Aldosterone
Adrenal gland
Angiotensin II
Angiotensinogen
Renin
production
Renin
Arteriole
constriction
Distaltubule
JGA
(b) The renin-angiotensin-aldosterone system (RAAS) leads to an increase
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 itskidneys offload much of the water absorbed froma 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 animals habitat
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Exploring environmental adaptations of the
vertebrate kidney
Fi 44 18
MAMMALS
Bannertail Kangaroo rat
(Dipodomys spectabi l is)
Beaver (Castor canadensis)
FRESHWATER FISHES AND AMPHIBIANS
Rainbow trout
(Oncorrhynchus mykiss)
BIRDS AND OTHER REPTILES
Roadrunner
(Geococcyx cal i fornianus)
Desert iguana
(Dipsosaurus dorsal is)
MARINE BONY FISHES
Northern bluefin tuna (Thunnus thynnus)