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CHAPTER
10
Ion and Water Balance
PowerPoint® Lecture Slides prepared by
Stephen Gehnrich, Salisbury University
Copyright © 2008 Pearson Education, Inc., publishing as Pearson Benjamin Cummings
Overview
ƒ Three homeostatic processes
ƒ Osmotic regulation
ƒ Osmotic pressure of body fluids
ƒ Ionic regulation
ƒ Concentrations of specific ions
ƒ Nitrogen excretion
ƒ Excretion of end-products of protein metabolism
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Ionic and Osmotic Challenges
ƒ Marine environments
ƒ Animals tend to gain salts and lose water
ƒ Freshwater environments
ƒ Animals tend to lose salts and gain water
ƒ Terrestrial environments
ƒ Animals tend to lose water
ƒ Many animals move between environments and
must be able to alter their homeostatic mechanisms
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Ionic Regulation
ƒ Strategies to meet ionic challenges
ƒ Ionoconformer
ƒ Ionoregulator
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Osmotic Regulation
ƒ Strategies to meet osmotic challenges
ƒ Osmoconformer
ƒ Internal and external osmolarity similar
ƒ For example, marine invertebrates
ƒ Omoregulator
ƒ Osmolarity constant regardless of external environment
ƒ For example, most vertebrates
ƒ Ability to cope with external salinities
ƒ Stenohaline
ƒ Can tolerate only narrow range
ƒ Euryhaline
ƒ Can tolerate wide range
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Osmolarity (mol/L): the osmolar concentration of a solution (commonly used in biology
Osmolality (mol/kg): the osmolal concentration of a solution
Osmotic concentration (Osm)
Salinity (%0, ppt, parts per thousand)
NaCl 150 mmol
Dissolve to Na+ 150 mmol, Cl- 150 mmol
Osmolarity = 300 mmol/L
Osmotic concentration = 300 mOsm
Water 1L
NaCl 500 mmol
Osmolarity = 1000 mmol/L
Osmotic concentration = 1000 mOsm
Salinity = 500*58 =29 g/L (ppt)
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Osmotic properties of cells
extracellular ion concentration
= intracellular ion concentation ?
Body fluid (extracellular) osmolarity
= intracellular osmolarity ?
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Osmoregulation in fishes
(cyclostome)
(freshwater fishes)
(Euryhaline fishes)
(seawater fishes)
freshwater
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seawater
Ionic and Osmotic Regulation
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Table 10.1
4 types of strategies for facing salt and water problems
Osmoconformer
Ionconformer
Osmoconformer
Ionregulator
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Osmoregulator Osmoregulator
Ionregulator
Ionregulator
Classification of Solutes
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Figure 10.4
Osmoconformers (but not ion-conformer)
The cells of osmoconformers are able to cope with high extracellar
Osmolarity by increasing their intracellular osmolarity.
Organic osmolytes: urea, trimethylamine oxide (TMAO)
Elasmobranch: shark, ray
hagfish
coelacanth
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crab-eating frog
Marine elasmobranch are hyperosmotic but hypoionic to seawater
Shark rectal gland
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Most types of FW animals share similar regulatory mechanism
Teleost
Ray (elasmobranches)
Lamprey (cyclostomes)
Frog
Soft-shell turtle
Mussel
Crayfish
Leech
Mosquito larvae
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Strategies of FW fishes:
Possessing an integument with a low permeability to salts and water
Do not drink water
Production of dilute urine
Reabsorption of salts from kidney
Ingesting salts from food
Active absorption of salts from skin (amphibian) or gills (fish)
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Osmoregulation in marine animals
Marine teleosts
Strategies:
Possessing an integument with a low
permeability to salts and water
Drink seawater
Production of isotonic urine
Excretion of salts from kidney (Mg2+,
SO42-)
Active secretion of salts from gills (Na+,
Cl-)
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Air-breathing animals: sea bird, sea turtle, iguanas,
Osmoregulatory problems:
Dehydration through their respiratory epithelia
Strategies:
Drink seawater
Production of isotonic urine
Active secretion of salts from salt glands (Na+, Cl-)
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Salt gland in sea birds (nasal gland)
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Shark rectal gland
Salt gland of
Sea turtle
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Salt gland of
marine Iguanas
Ion- and osmo-regulation of animals:
from molecular to cellular function
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Relative permeability of phosolipid bilayer (cell membrane) to
molecules and ions
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Mechanisms for trans-membrane movement of ions
Diffusion (move down electrochemical gradient)
Passive transport (move down electrochemical gradient)
Active transport (move against electrochemical gradient)
Passive transport:
Facilitated
Ion channels
Carrier proteins diffusion
Active transport:
Primary active transport
Secondary active transport
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5 types of ion transporters
Primary active transport
Passive transport
secondary active transport
Passive transport
cotransporter
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exchanger
Passive transport through “passive transporter”
Facilitated diffusion
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Primary active transport through Na+/K+-ATPase (Na+ pump)
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3 major types of active transporters:
Na pump (Na/K-ATPase)
Ca pump (Ca-ATPase)
H pump (H-ATPase,
V-ATPase)
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Secondary active transportrs
Na/H exchanger
Na/Ca exchanger
Na/K/Cl cotransporter
Cl/HCO3 exchanger
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Kinetics of various ion transport proteins
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Epithelial Tissue
ƒ Epithelial tissues form boundary between animal
and environment
ƒ External surfaces
ƒ For example, skin
ƒ Internalized surfaces
ƒ For example, lumen of digestive and excretory systems
ƒ Epithelial tissues have physiological functions in
respiration, digestion, and ion and water regulation
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Epithelial Tissue Properties for
Ion Movement
ƒ Four features of transport epithelia
ƒ Asymmetrical distribution of membrane transporters
ƒ Solutes selectively transported across membrane
ƒ Cells interconnected to form impermeable sheet of
tissue
ƒ Little leakage between cells
ƒ High cell diversity within tissue
ƒ Abundant mitochondria
ƒ Large energy (ATP) supply
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Epithelial Tissue Properties for
Ion Movement
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Figure 10.8
Solute Movement
ƒ Epithelial cells use two main routes of transport
ƒ Transcellular transport
ƒ Movement through the cell across membranes
ƒ Paracellular transport
ƒ Movement between cells
ƒ “Leaky” vs. “tight” epithelia
ƒ Types of transporters
ƒ Na+/K+ATPase
ƒ Ion channels (Cl–, K+, Na+)
ƒ Electroneutral cotransporters
ƒ Electroneutral exchangers
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Transcellular and Paracellular Transport
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Figure 10.9
Measurement of trans-epithelial ion transport
Ussing chamber
Voltage/ Ion clamp technique
(Short Circuit Current Vs = 0)
ClTransepithelial potential Vout = 0
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Vin = -30 mV
Epithelial Cells in Fish Gills
ƒ Fish gill lamellae composed of
ƒ Mitochondria-rich chloride cells
ƒ Pavement cells
ƒ Some mitochondria-rich ??
ƒ Some mitochondria-poor
ƒ Transport likely carried out by mitochondria-rich cells
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Salt secretion in marine teleosts
Salt-secreting cells (chloride cells)
Gill filament
Branchial chloride cells in gill filament of marine teleosts
lamellae
filament
Salt-secreting cells present in :
rectal gland of shark, sea bird, sea turtle
gills of marine teleosts
K channel
Na/K ATPase
Epithelial Cl channel
(CFTR)
Paracellular
pathway
Na/K/2Cl
cotransporter
Leaky junction
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Epithelial Cells in Fish Gills (FW fish)
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Figure 10.10
Ion Transport by Fish Gills
Direction of ion transport depends on water salinity
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Figure 10.11
Hypothesis of Na, Cl uptake in freshwater fish
Gas exchange
Ion exchange
pH regulation
Ammonia excretion
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Renal physiology
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The Kidney
ƒ Vertebrate kidneys have six roles in homeostasis
ƒ Ion balance
ƒ Osmotic balance
ƒ Blood pressure
ƒ pH balance
ƒ Excretion of metabolic wastes and toxins
ƒ Hormone production
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Kidney Structure and Function
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Figure 10.19
The Nephron
ƒ Functional unit of the kidney
ƒ Composed of
ƒ Renal tubule
ƒ Lined with transport epithelium
ƒ Various segments with specific transport functions
ƒ Vasculature
ƒ Glomerulus
ƒ Ball of capillaries
ƒ Surrounded by Bowman’s capsule
ƒ Capillary beds surrounding renal tubule
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Structure of nephron
15%
Glomerulus
Renal corpuscle
Bowman’s capsule
Proximal convoluted tubule
Proximal tubule
Proximal straight tubule
Descending thin limb of Henle’s loop
Henle’s loop
Ascending thin limb of Henle’s loop
Thick ascending limb of Henle’s loop
Distal tubule
Collecting duct
Distal convoluted tubule
Cortical collecting duct
Medullary collecting duct
Renal pelvis
Ureter
Bladder
Urethra
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Urine Production
ƒ Four processes
ƒ Filtration
ƒ Filtrate of blood formed at glomerulus
ƒ Reabsorption
ƒ Specific molecules in the filtrate removed
ƒ Secretion
ƒ Specific molecules added to the filtrate
ƒ Excretion
ƒ Urine is excreted from the body
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Filtration
ƒ Liquid components of the blood are filtered into
Bowman’s capsule
ƒ Water and small solutes cross glomerular wall
ƒ Blood cells and large macromolecules are not filtered
ƒ Glomerular capillaries are very leaky
ƒ Podocytes with foot processes form filtration structure
ƒ Mesangial cells control blood pressure and
filtration within glomerulus
ƒ Filtrate flows from Bowman’s capsule into
proximal tubule
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Blood pressure in the renal glomerulus
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Net filtration pressure
GFR
Permeability of bowman’s
capsule
blood
A B C
urine
blood
protein
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A: endothelium (pores)
B: basement membrane
C: podocyte (filtration slit)
Intrinsic control of GFR:
1. Autoregulation of blood pressure of afferent arteriole
2. Renal blood flow regulated by juxtaglomerular apparatus
(macula densa, juxtaglomerular cells)
3. Sympathetic activation causes vasocontriction of afferent arteriole
and reduces hydraulic permeability (podocytes) of Bowman’s
capsule
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Extrinsic Regulators of GFR
ƒ Hormones
ƒ Vasopressin (antidiuretic hormone, ADH)
ƒ Renin-Angiotensin-Aldosterone (RAA) pathway
ƒ Atrial natriuretic peptide (ANP)
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Reabsorption
ƒ Primary urine
ƒ Initial filtrate filtered in Bowman’s capsule that is
isosmotic to blood
ƒ Most water and salt in primary urine reabsorbed
using transport proteins and energy
ƒ Rate of reabsorption limited by number of transporters
ƒ Renal threshold
ƒ Concentration of a specific solute that will overwhelm
reabsorptive capacity
ƒ Each zone of the nephron has transporters for
specific solutes
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Reabsorption of Glucose
Glucose is reabsorbed by secondary active transport
Reabsorbed molecules taken up by the blood
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Figure 10.23
Renal clearance of substance =
Amount of substance in urine
Amount of substance filtered
Glucose clearance= 0
(100% reabsorption)
Inulin clearance=1
(no reabsorption, secretion)
GFR (L/h) =
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Inulin in urine (mg/h)
Inulin in blood (mg/L)
Transport in Tubule Regions
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Figure 10.25
Transport in the Proximal Tubule
ƒ Most reabsorption of solutes and water takes place
in proximal tubule
ƒ Many solutes reabsorbed by Na+ cotransport
ƒ Water follows by osmosis
ƒ Proximal tubule also carries out secretion
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Figure 10.27
Secretion
ƒ Similar to reabsorption, but in reverse
ƒ Molecules removed from blood and transported into
the filtrate
ƒ Molecules secreted include
ƒ K+, NH4+, H+, pharmaceuticals, and water-soluble
vitamins
ƒ Requires transport proteins and energy
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Ion and Water Transport in the Loop of Henle
ƒ Descending limb is permeable to water
ƒ Water is reabsorbed
ƒ Volume of primary urine decreases
ƒ Primary urine becomes more concentrated
ƒ Ascending limb is impermeable to water
ƒ Ions are reabsorbed
ƒ Primary urine becomes dilute
ƒ Reabsorbed ions accumulate in interstitial fluid
ƒ An osmotic gradient created in the medulla
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Urine concentration: Henle’s loop
Reabsorption
of water 70%
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Reabsorption
of water 30%
(regulatory region)
Primary active Na reabsorption
Coupling of water reabsorption to Na reabsorption
Osmolarity
increase
water
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Proximal tubule:
Na reabsorption
Na/glucose cotransporter
Na/K/Cl cotransporter
Ascending limb of Henle’s loop:
Na, reabsorption
Na/H exchanger
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Urine concentration: countercurrent multiplier system
Renal cortex
low
Interstitial
osmolarity
high
Renal medulla
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Countercurrent flow of circulation (vasa recta)
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Renal Na and water regulation:
Control of GFR (short-term)
Control of Na reabsorption (long-term)
Renin-angiotensin
Aldosterone
hormones
baroreceptor
Atrial natriuretic peptide
(ANP)
Vasopressin
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Renin-angiotensin system and aldosterone
Activity of renal
sympathetic nerve
Intrarenal
Baroreceptor
(JGA)
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Renin-angiotensin system
Angiotensin-converting
enzyme (ACE)
Aldosterone (mineral corticoid)
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Atrial natriuretic peptide (ANP) stimulates
Na excretion
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Vasopressin
ƒ Also called antidiuretic hormone (ADH)
ƒ Peptide hormone
ƒ Produced in hypothalamus and released by posterior
pituitary gland
ƒ Increases water reabsorption from the collecting duct
by increasing number of aquaporins
ƒ Release stimulated by increasing plasma osmolarity
detected by osmoreceptors in the hypothalamus
ƒ Release is inhibited by increasing blood pressure
detected by stretch receptors in atria and baroreceptors
in carotid and aortic bodies
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Regulation of water reabsorption
Collecting duct
Aquaporin (water channel)
Vasopressin regulated
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Vasopressin Increases Cell Permeability
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Figure 10.34a
Renal (metabolic) acid-base regulation in distal tubule and
collecting duct
A-type intercalated cell
Acid secretion
Apical H-ATPase
(proton-ATPase)
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B-type intercalated cell
Base (bicarbonate) secretion
Apical Cl/bicarbonate exchanger
(anion exchanger)