Ion and Water Balance

Part 1

Chapter 10; 08.11.-12.11.2010

Overview
Environment has several different meanings
External world for the whole animal
Extracellular fluid for a cell
Cytoplasm for intracellular enzymes

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Overview

Mud puppy (Necturus maculosus)

Animals use different

combinations of
tissues to control ion
and water balance in
various internal and
external environments
Water and ion exchanges
Between:
External fluid
Internal fluids:
interstitial fluid
extracellular fluid
plasma
intracellular fluid
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Figure 10.1

Overview
Three homeostatic processes
Osmotic regulation
Osmotic pressure of body fluids

Ionic regulation
Concentrations of specific ions

Nitrogen excretion
Excretion of end-products of protein metabolism

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Overview
Three homeostatic processes
Osmotic regulation is the control of tissue osmotic
pressure, which determines the driving force for the
movement of of water across biological membranes.
Animals and cells cannot actively pump water.
Osmotic regulation requires the movement of solutes
across membranes, altering osmotic gradients.

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Overview
Three homeostatic processes
Ionic regulation is the control of the ionic composition
of body fluids (well focus on the ions that are
important solutes, and therefore relevant to the
osmoregulatory strategies).

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Overview
Three homeostatic processes
Nitrogen excretion is the pathway by which animals
excrete ammonia, the toxic nitrogenous end product of
protein catabolism. The process for expelling
ammonia, or metabolic alternatives such as urea and
uric acid, is linked to the control of osmotic and ionic
homeostasis.

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Ionic and Osmotic Challenges

Marine environments
Animals tend to gain salts and lose water

Freshwater environments
Animals tend to lose salts and gain water

Terrestrial environments
Animals tend to lose water

Many animals move between environments and

must be able to alter their homeostatic mechanisms

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ALL multicellular animals regulate ICF

composition of ECF and ICF differ

OP similar
Plasma membrane maintains ionic, but not osmotic,
difference between ICF and ECF
Do animals regulate ECF?

For animals that live in

water:
does OPECF = OPext env?
does [ion]ECF = [ion]ext env?

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Ionic and Osmotic Challenges

Ionoregulatory and osmoregulatory strategies of
aquatic animals can be distinguished by:
1. the differences between extracellular fluids and
external conditions;
2. the extent to which extracellular fluids change
when external conditions change.

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Ionic and Osmotic Regulation

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Table 10.1

Ionic Regulation
Strategies to meet ionic challenges
Ionoconformer
Exert little control over ion profile within the extracellular
space
Exclusively found in marine animals
For example, many invertebrates

Ionoregulator
Control ion profile of extracellular space
For example, most vertebrates

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Osmotic Regulation
Strategies to meet osmotic challenges
Osmoconformer
Internal and external osmolarity similar
For example, marine invertebrates

Omoregulator
Osmolarity constant regardless of external environment
For example, most vertebrates

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Ionic and Osmotic Regulation

Ability to cope with external salinities
Stenohaline
Can tolerate only narrow range

Euryhaline
Can tolerate wide range

NB: There is no predetermined relationship between the

strategy (osmoconforming versus osmoregulating) and the
degree of tolerance (euryhaline versus stenohaline) - see
Fig. 10.2.

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Ionic and Osmotic Regulation

Osmotic strategies can be distinguished by three factors: 1. the osmotic

gradients between the animal and the water; 2. the degree to which internal
osmolarity changes in relation to a changing external osmolarity; 3. the degree
of tolerance of an osmotic challenge.
The four osmotic strategies depicted in this figure can be distinguished by
following the internal osmolarity of four animals living in full-strength
seawater, then exposed to a decreasing osmolarity until death.
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Stenohaline
Can tolerate only
narrow range
Euryhaline
Can tolerate wide
range

Figure 10.2

Osmoregulatory strategies (and salinity tolerance)

Fig. 11.2

Stenohaline
Can tolerate only narrow range
Euryhaline
Can tolerate wide range

Sources of Water and Solutes

The environment provides water in many forms:
Dietary water
Water preformed in plant and animal tissue

Metabolic water
Water generated as result of oxidative phosphorylation

Drinking

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Dietary Water

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Table 10.2

Classification of Solutes
Three classes of solutes are distinguished by their effects
on the structure and function of macromolecules, such as
enzymes:
Perturbing
Disrupt macromolecular function at normal concentrations found within
the animal
Na+, K+, Cl, SO4+, charged amino acids

Compatible
Little affect on macromolecular function and can accumulate to high
concentration without deleterious effects on cellular processes
Polyols (glycerol, glucose) and uncharged amino acids

Counteracting
Disrupt function on their own, but counteract disruptive effects of other
solutes when employed in combination. Urea/Methylamines (betain)
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Classification of Solutes

Each type of solute exerts characteristic effects on macromolecular structure and

function, such as enzyme kynetics.
a. A perturbing solute is shown to increase the Km value of a hypothetical
enzyme, whereas a compatible solute at the same concentration has no effect
on Km.
b. Each counteracting solute has perturbing effects when present alone, but when
both both are present, the effects are offset.
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Figure 10.3

Classification of Solutes

Figure 10.4
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The relative importance of the

various solutes in different
species and cellular
compartments.
Seawater is mainly Na and
Cl, with lower levels of
other ions, such as K,
Mg2+, and Ca2+.
Ionoconformers have high
levels of Na and Cl,
whereas the levels of these
ions are lower in
ionoregulators.
Osmoconformers have the
same osmolarity as seawater
but maintain an inorganic
ion profile much like that of
an osmoregulator. The
remainder of the osmolarity
is due to organic solutes,
such as urea, amino acids,
and methylamines.

Cell Volume
Cells transport solutes in and out of extracellular
fluid (ECF) to control cell volume
Water follows solutes by osmosis

Animal regulates composition of the ECF

Provides cells with external solution that allows them
to maintain appropriate cell volume

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Regulation of Cell Volume

Cells actively change cell
volume by moving ions in or
out of the cell. Each of the
cells depicts a combination
of transporters commonly
used to transfer ions into the
cell (causing a volume
increase) or out of the cell
(causing cell shrinking).

Figure 10.5
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Epithelial Tissue
The role of epithelial tissue
Epithelial tissues form boundary between animal
and environment
External surfaces
For example, skin

Internalized surfaces
For example, lumen of digestive and excretory systems

Epithelial tissues have physiological functions in

respiration, digestion, and ion and water regulation

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Integument
Animals change flux of water across body surface
by mediating permeability of the integument
Aquaporin proteins increase water permeability 100fold

Typically animals need to reduce water flux

Cover external surfaces with layer of hydrophobic
molecules
Mucus
Cornified stratum corneum with keratin
For example, terrestrial amniotes
Cuticle with chitin
For example, arthropods
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Integument

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Figure 10.6

Integument

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Figure 10.7

Epithelial Tissue Properties for

Ion Movement
Four features of transport epithelia
Asymmetrical distribution of membrane transporters
Solutes selectively transported across membrane

Cells interconnected to form impermeable sheet of

tissue
Little leakage between cells

High cell diversity within tissue

Abundant mitochondria
Large energy (ATP) supply

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Epithelial Tissue Properties for

Ion Movement

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Figure 10.8

Solute Movement
Epithelial cells use two main routes of transport
Transcellular transport
Movement through the cell across membranes

Paracellular transport
Movement between cells
Leaky vs. tight epithelia

Types of transporters
Na+/K+ATPase
Ion channels (Cl, K+, Na+)
Electroneutral cotransporters
Electroneutral exchangers
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Transcellular and Paracellular Transport

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Figure 10.9

Epithelial Cells in Fish Gills

Fish gill lamellae composed of
Mitochondria-rich chloride cells
Pavement cells
Some mitochondria-rich
Some mitochondria-poor

Transport likely carried out by mitochondria-rich cells

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Epithelial Cells in Fish Gills

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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

Saltwater-Freshwater Transitions
Some fish migrate between saltwater and
freshwater (diadromous fish)
Catadromous
Live in freshwater, migrate to saltwater to spawn
For example, eels
Live in saltwater; migrate to freshwater to spawn
For example, salmon

Ion transport functions of epithelia must change

during migration
Controlled by hormones

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Saltwater-Freshwater Transitions

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Figure 10.12

Digestive Epithelia
Water and salts from drinking and food transported
across digestive epithelium
Transcellular and paracellular transport involved
Absorbed water and salts enter blood

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Salt Glands
Reptiles and birds possess salt glands
Glands located near the eye, drain into ducts that
empty near nostril
Excrete hyperosmotic solutions of Na+ and Cl
Large amount of salt excreted in small volume of water

Hyperosmotic solutions produced by ion pumps and a

countercurrent multiplier

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Salt Glands

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Figure 10.13

Rectal Glands
Accessory excretory organ in elasmobranchs
Empties into digestive tract

Actively transport Na+ and Cl from blood into

lumen of the gland
Ion transport similar to chloride cell and salt glands
Rate of salt excretion regulated by hormones

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Rectal Glands

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Figure 10.14

Nitrogen Excretion
Ammonia produced during amino acid breakdown
is toxic and must be excreted
Ammonia nitrogen excreted in three forms
Ammonia (ammonioteles)
Uric acid (uricoteles)
Urea (ureoteles)

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Figure 10.15

Nitrogen Excretion
Type of nitrogen
compound excreted is
related to the animals
environment
Aquatic animals usually
excrete ammonia
Terrestrial animals usually
excrete urea or uric acid
Many animals change
mode of nitrogen
excretion in response to
water availability
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Table 10.3

Ammonia Excretion
Advantages
Ammonia released by
deamination of amino
acids
Requires little energy to
produce

Disadvantages
Highly toxic
Requires large volumes
of water to store and
excrete
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Figure 10.16

Uric Acid Excretion

Advantages
Few toxic effects
Can be excreted in
small volume of water

Disadvantages
Expensive to produce

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Figure 10.17

Urea Excretion
Advantages
Only slightly toxic
Relatively inexpensive to produce

Disadvantages
Urea is a perturbing solute

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Figure 10.18

Urea as an Osmolyte
Cartilaginous fish use urea to increase tissue
osmolarity
Helps prevent water loss in marine environment
Ureas perturbing effects counteracted by
methylamines (TMAO, betaine, sarcosine)

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