Chapter 44

Osmoregulation and Excretion
PowerPoint Lectures for Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero
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• 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 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

Figure 44.3a
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(a) Osmoregulation in a saltwater fish

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
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
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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 (2 mL/day = 100%) Ingested in food (0.2) Water gain Ingested in liquid (1,500) Water balance in a human (2,500 mL/day = 100%) Ingested in food (750)

Derived from metabolism (1.8)

Derived from metabolism (250)

Feces (0.9) Urine (0.45) Urine (1,500)

Feces (100)

Water loss

Figure 44.5
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Evaporation (1.46)

Evaporation (900)

• 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%.

Water lost per day (L/100 kg body mass)

4 3 2 1 0 Control group (Unclipped fur) Experimental group (Clipped fur)

CONCLUSION

The fur of camels plays a critical role in their conserving water in the hot desert 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 blood
Nasal salt gland (a) An albatross’s 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. Nostril with salt secretions Lumen of secretory tubule Vein Capillary Secretory tubule Transport epithelium Direction of salt movement Artery
NaCl

(b) One of several thousand secretory tubules in a saltexcreting gland. Each tubule is lined by a transport epithelium surrounded by capillaries, and drains into a central duct.

Blood Secretory cell flow of transport epithelium Central duct

(c) The secretory cells actively transport salt from the blood into the tubules. Blood flows counter to the flow 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.

Figure 44.7a, b

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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 Amino acids Nucleic 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 HN O C C H C N C O

NH3

O

C

NH2 NH2 Urea

C N N H H Uric acid

Figure 44.8

Ammonia

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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 Excretory tubule Filtrate Urine 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.

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

Figure 44.10
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Nephridiopore in body wall

• 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

Nephridiopore

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

Malpighian tubule Rectum

Figure 44.12

HEMOLYMPH

Reabsorption of H2O, ions, and valuable organic molecules

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

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 (b) Kidney structure

Figure 44.13b

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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
JuxtaCortical medullary nephron nephron
Afferent arteriole from renal artery Glomerulus Bowman’s capsule Proximal tubule Peritubular capillaries Collecting duct SEM Efferent arteriole from glomerulus Branch of renal vein Descending limb Ascending limb Vasa recta Distal tubule Collecting duct

Renal cortex

20 µm Renal medulla

To renal pelvis

Loop of Henle

Figure 44.13c, d
(c) Nephron Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (d) Filtrate and blood flow

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 NaCl Nutrients HCO3− H2 O K+ 4 Distal tubule NaCl H2 O HCO3−

H+

NH3

K+

H+

CORTEX Filtrate H2O Salts (NaCl and others) HCO3– H+ Urea Glucose; amino acids Some drugs 2 Descending limb of loop of Henle H2 O OUTER MEDULLA 3 Thin segment of ascending limb NaCl 5 Collecting duct Urea NaCl INNER MEDULLA H2 O 3 Thick segment of ascending limb NaCl

Key Active transport Passive transport

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
300 300 300 100 300 CORTEX Active transport Passive transport H2 O H2 O H2 O H2 O H2 O H2 O INNER MEDULLA H2 O 400 NaCl NaCl NaCl NaCl NaCl NaCl NaCl 200 H2 O H2 O H2 O H2 O H2 O 700 Urea H2 O Urea H2 O Urea 1200 1200 900 400 400 300 100 Osmolarity of interstitial fluid (mosm/L)

OUTER MEDULLA

600

400

600

600

900

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 Hypothalamus Drinking reduces blood osmolarity to set point ADH Increased permeability Pituitary gland Distal tubule Thirst

STIMULUS: The release of ADH is triggered when osmoreceptor cells in the hypothalamus detect an increase in the osmolarity of the blood

H2O reabsorption helps prevent further osmolarity increase Collecting duct

Homeostasis: Blood osmolarity

Figure 44.16a

(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 functions in homeostasis
Increased Na+ and H2O reabsorption in distal tubules Homeostasis: Blood pressure, volume

Aldosterone

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

Adrenal gland

Angiotensin II Distal tubule Angiotensinogen JGA Renin production Renin

Figure 44.16b

(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 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)

Beaver (Castor canadensis) FRESHWATER FISHES AND AMPHIBIANS

Desert iguana (Dipsosaurus dorsalis) MARINE BONY FISHES

Rainbow trout (Oncorrhynchus mykiss)

Northern bluefin tuna (Thunnus thynnus)

Figure 44.18

Frog (Rana temporaria)

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