Comparative Vertebrate Anatomy

Lecture 10

Living in Water
(Respiration, buoyancy, sensation, and water balance
in aquatic vertebrates)
 The Aquatic Environment

Physical properties of water create some difficulties for aquatic

animals
Buoyancy in water column
Water is a dense medium to move through
Heat flows rapidly between an animal and the water around it
Ions and water molecules move readily between the external
environment and an animals body fluids maintaining
homeostasis is difficult (but disposal of nitrogenous wastes is
easier)
Concentration of oxygen is lower than it is in air, and density of
water imposes limits on kinds of gas exchange structures that
can be effective
 The Aquatic Environment

73% of Earths surface is covered by water, only about 0.01% of this

water is contained in freshwater lakes, rivers, etc.
However, freshwater systems are extremely rich biologically, and
nearly 40% of all bony fishes live in freshwater
Water and air provide drastically different environments for
vertebrates
E.g., gravity is an important force acting on a terrestrial animal,
but air resistance is trivial for all but the fastest birds
In water, gravity is negligible, but fluid resistance to movement is
a major factor
The Aquatic Environment
 Gills Obtaining oxygen in water

Gills are specialized structures for oxygen/carbon dioxide exchange

Gills of teleosts (derived ray-finned fishes that include the majority of
extant freshwater and marine fishes) are enclosed in opercular
cavities
Flow of water is usually unidirectionalin through mouth, out
through gills
Flaps inside the mouth and at the margins of the gill covers
(opercula) act as valves to prevent backflow
Respiratory surfaces are delicate projections from the lateral side of
each gill arch two columns of gill filaments extend from each gill
arch
As water leaves the buccal cavity, it passes over the filaments
and gas exchange takes place at the numerous microscopic
projections from the filaments called secondary lamellae
Gills Obtaining oxygen in water
Gills Obtaining oxygen in water
 Gills Obtaining oxygen in water

Pumping action of the mouth and opercular cavities (buccal

pumping) creates positive pressure across the gills
Some filter-feeding and pelagic fishes (mackerel, tunas,
swordfishes, some sharks) have reduced or lost the ability to pump
water across the gills
These fishes must create a respiratory current by ram
ventilation
 Gills Obtaining oxygen in water

Arrangement of blood vessels in the gills maximizes oxygen

exchange
Each gill filament has two arteries, an afferent vessel running
from the gill arch to the filament and an efferent vessel returning
blood to the arch
Each secondary lamellae is a blood space connecting the
afferent and efferent vessels
Direction of blood flow through the lamellae is opposite to the
direction of water flow across the gill countercurrent exchange
Maximizes oxygen content that can diffuse into blood
Countercurrent flow maintains
a diffusion gradient between
blood and water for the full
length of the lamella, and
results in high oxygen
concentration in the blood
leaving the gills
 Lungs and other Respiratory Structures

Fishes in environments with low oxygen levels supplement oxygen

from gills with additional oxygen obtained from the air via lungs or
accessory air respiratory structures (gulp air)
Tropical anabantid fishes suck air into the mouth and transfer it
to vascularized chambers in the rear of the head, called
labyrinths
Many of these fishes are facultative breathers, but some (electric
eels and some snakeheads) are obligatory air breathers (gills alone
cannot meet respiratory demands)
Lungs first appeared in fishes, develop embryonically as
envaginations of the pharyngeal region of the digestive tract
Lungs of bichirs, lungfishes, and tetrapods originate from the
ventral surface of the gut, whereas lungs of gars and teleosts
originate from dorsal surface
 Adjusting Buoyancy

Altering volume of air in lungs or bladders can allow aquatic

vertebrates to adjust buoyancy
Many teleosts are neutrally buoyant and do not have to swim to
maintain vertical position in the water column (have well developed
swim bladders)
Only movements necessary are backpedaling of the pectoral
fins to counteract the thrust produced by water ejected from the
gills
The swim bladder is located between the peritoneal cavity and the
vertebral column
Occupies ~5% of body volume in marine teleosts and ~7% in
freshwater teleosts (salt water is denser)
The bladder wall has smooth walls of interwoven collagen fibers
without blood vessels
To maintain buoyancy, a fish must adjust the volume of gas in its
swim bladder as it changes depth
 Adjusting Buoyancy

Volume is regulated by secreting gas into the bladder to counteract

the increased external water pressure as the fish descends, and by
removing gas when it ascends
Basal teleosts (e.g., goldfish, eels, salmon, minnows) are
physostomous and retain a connection (pneumatic duct) between
the gut and swim bladder
Can gulp air at the surface to fill the bladder and burp gas to
reduce its volume
Pneumatic duct is absent from more derived teleosts, a condition
termed physoclistus, and these fish secrete gas from the blood into
the bladder
Both physostomes and physoclists have a gas gland located in
the anterior ventral floor of the bladder
Underlying the gland are many capillaries arranged to give
countercurrent flow of blood entering and leaving the area a
structure called the rete mirabile (wonderful net)
 Adjusting Buoyancy

The rete mirabile is effective at extracting gas (especially oxygen)

from the blood and releasing it into the swim bladder
The gas gland secretes oxygen by releasing lactic acid and CO2,
which acidifies the blood in the RM
Acidification causes hemoglobin to release oxygen into solution
Oxygen released accumulates in the rete until its pressure
exceeds the oxygen pressure of the swim bladder, at which point
oxygen diffuses into the bladder (possible because of
countercurrent position of capillaries in rete and the Root
effect)??
To release excess gas, physoclists open a muscular valve, the
ovale, adjacent to a capillary bed
High internal pressure of oxygen causes it to diffuse into the
blood when the ovale sphincter is opened
Adjusting Buoyancy
 Adjusting Buoyancy

The Bohr effect is a reduction in the

affinity of hemoglobin for oxygen in
the presence of acid hemoglobin
releases oxygen

The Root effect is a reduction in the

maximum amount of oxygen that
hemoglobin can bind
 Adjusting Buoyancy

Some deep sea teleosts have lost the swim bladder

Blobfishes (Psychrolutes marcidus) occur at depths of ~1000 m
Flesh is a gelatinous mass with very little muscledensity
slightly less than water
Floats above sea floor without expending energy on buoyancy
 Adjusting Buoyancy

Cartilaginous fishes do not have swim bladders; instead use the

liver to create neutral buoyancy
The liver of a shark has high oil content, making sharks lower
density (lighter) than sea water
Bottom-dwelling sharks have livers with fewer and smaller oil
vacuoles in their cells, and are negatively buoyant
Nitrogen-containing compounds in blood of cartilaginous fishes also
contributes to their buoyancy
 Water and the Sensory World of Fishes

Because water absorbs light, and light is scattered by suspended

particles, objects become invisible at a distance of a few hundred
meters in even the clearest water
Fishes supplement vision with other senses, some of which operate
only in water (e.g., detecting water movement via lateral line system,
electrical sensitivity)
In terrestrial vertebrates cornea plays a major role in focusing an
image, whereas in aquatic vertebrates the lens plays a major role in
focusing light on the retina
Aquatic vertebrates have spherical lenses with high refractive
indices
 Chemosensation: Taste and Odor

Fish have taste-bud organs in the

mouth and around the head and
anterior fins olfactory organs on
the snout detect soluble
substances
sharks and salmon can detect
odors at concentrations of less
than 1 part per billion
Homeward-migrating salmon
are directed to their stream of
origin from large distances by
a chemical signature from the
home stream that was
imprinted when they were
juveniles
 Lateral Line System Mechanical Reception

Mechanical receptors detect touch, sounds, pressure, and motion

Like all vertebrates, fishes have an internal ear (labyrinth organ)
that detects changes in speed and direction of motion also
gravity detectors at the base of semicircular canals to distinguish
up from down
Most also have an auditory region sensitive to sound-pressure
waves
In fishes and aquatic amphibians, clusters of hair cells and support
cells form neuromast organs that are dispersed over the head
and body
In jawed fishes, neuromast organs are located in a series of
canals on the head, and one or more canals pass along the
sides of the body onto the tail
This surface receptor system is the lateral line system
(detects water displacement)
 Lateral Line System Mechanical Reception

Neuromasts are distributed either in tubular canals or exposed in

epidermal depressions (some fishes have both arrangements)
Hair cells have a kinocilium placed asymmetrically in a cluster of
smaller hairs (microvilli)
Cells are arranged in pairs with the kinocilia positioned on
opposite sides of adjacent cells
A neuromast contains many such hair-cell pairs
Each neuromast has two afferent nerves one transmits
impulses from cells with kinocilia in one direction, the other for
cells in the opposite direction
Kinocilia and microvilli are embedded in a gelatinous secretion,
the cupula displacement causes kinocilia to bend
Resultant deformation either excites or inhibits a neuromasts
nerve discharge (depending on direction)
Net effects of cupula displacement is to increase firing rate in
one afferent nerve and to decrease it in the other nerve
 Lateral Line System Mechanical Reception

E.g., killifish detect insects on the water surface with lateral line
system each neuromast group on the head provides information
about surface waves coming from a different direction (with
overlapping stimulus fields)
 Electrical Discharge and Reception

Unlike air, water conducts electricity, and some fishes can discharge
enough electricity to stun prey animals and deter predators
Other weakly electric fishes use electrical signals for courtship and
territorial defense
All use modified muscle tissue to produce electrical discharge
(modified muscle cells are called electrocytes have lost the
capacity to contract and are specialized for generating an ion current
flow)

Electric Catfish
 Electrical Discharge and Reception

When at rest, membranes of muscle cells and nerve cells are

electrically charged, with intracellular fluids about 84 millivolts more
negative than extracellular fluids (due to exclusion of sodium)
When cell is stimulated, sodium ions flow rapidly across the smooth
surface into the cell, sending its potential to +67 millivolts (difference
of 151 mV)
Electrocytes are arranged in stacks, like batteries in a flashlight, so
individual potentials are combined to produce high voltages
South American electric eel has up to 10,000 layers of cells and
can generate potentials in excess of 600 volts
Electrical Discharge and Reception
 Electrical Discharge and Reception

High conductivity of seawater makes it possible for sharks to detect

electrical activity generated by muscle contractions of their prey
Sharks and rays have sensitive electroreceptors called ampullae of
Lorenzini on their heads
Pores filled with electrically conductive gel, that can detect
changes in electrical potential in space
Sharks may also use electroreception for navigation
(electromagnetic field of Earth and ocean currents carrying ions
produce tiny electrical gradients)
Electrical Discharge and Reception
Electrical Discharge and Reception
Electrical Discharge and Reception
 Electrical Discharge and Reception

Some freshwater fishes produce weak electrical discharge (no direct

offensive or defensive value)
Most of these fishes are nocturnal and live in very turbid waters
These fishes use discharges for electrolocation and social
communications (in pulses)
Fish detects the presence, position, and movement of objects by
sensing where on its body maximum distortion of its electric field
occurs (distortion occurs by presence of electrically conductive [e.g.
other fish] or resistive [e.g., rocks] objects in the surroundings)
Electroreceptors are modified neuromast receptors
Different durations and frequencies of discharges distinguish
immature individuals, females with eggs, and sexually active
males in many species
 Electrical Discharge and Reception

A electric field surrounds a weakly electric fish electroreceptors

in the skin allow the fish to detect the presence of nearby objects
by sensing distortion in the electrical force
 Internal Environment of Vertebrates

70-80% of body mass of most vertebrates is water

Chemical reactions that release energy or synthesize new
chemical compounds take place in an aqueous environment
Body fluids of vertebrates contain a complex mixture of ions and
other solutes
Cofactors that control rates of metabolic processes
Regulate pH, stability of cell membranes, or electrical activity of
nerves
Metabolic substrates must diffuse from sites of synthesis to sites
of utilization
Almost everything that occurs in body tissues of vertebrates
involves water, and maintaining concentrations of water and
solutes within narrow limits is vital
 Internal Environment of Vertebrates

Freshwater vertebrates face the threat of being flooded with water

that flows into them from the environment
Saltwater vertebrates must prevent water in their bodies from being
sucked out into the sea
Temperature is also a critical factor for living organisms chemical
reactions are temperature sensitive
Water temperature is more stable than air temperature because
water has a much higher heat capacity simplifies the task of
maintaining a constant body temperature (as long as optimal body
temperature corresponds to water temperature)
Aquatic animals have a harder time maintaining a body
temperature different from water temperature because water
conducts heat so well
Internal Environment of Vertebrates
 Exchange of Water and Ions

All living animals contain organic and inorganic substances

dissolved in water, enclosed by a permeable body surface
Exchange of matter and energy with the environment is essential to
survival much of this exchange is regulated by the body surface
Water and ions move across quite freely, whereas larger
molecules do not (differential permeability)
Vertebrates use both active and passive exchange to regulate
internal conditions
The kidney plays a crucial role in controlling water balance and
excreting wastes
Organisms can tolerate only a narrow range of concentrations of
body fluids
Must eliminate waste products (especially ammonia) before they
reach harmful levels
 Exchange of Water and Ions

Adult vertebrate kidney consists of hundreds to millions of

nephrons, each of which produces urine
Primary function is to remove excess water, salts, waste
metabolites, and foreign substances from the blood
Blood is first filtered through a glomerulus, a structure unique to
vertebrates
Composed of a leaky arterial capillary tuft encapsulated within a
sieve-like filter
Arterial blood pressure forces fluid in the nephron to form an
ultrafiltrate, composed of blood minus the cells and larger
molecules
Ultrafiltrate is processed to return water and essential
metabolites to circulation
Remaining fluid is urine
 Exchange of Water and Ions

First vertebrates likely had ion levels similar to those in seawater

(e.g., hagfishes) like most marine invertebrates
Salt concentrations are greatly reduced in blood of all other
vertebrates
Salt ions, urea, and some small carbohydrate molecules are
solutes primarily involved in regulating body fluid concentrations
Water moves from areas of high water potential to areas of lower
potential (presence of solutes lowers potential activity of water)
Water flows from a dilute solution (high water potential) to a
more concentrated solution (low water potential) osmosis
 Exchange of Water and Ions

Seawater has a solute concentration of ~1000 mmol kg-1

Hagfishes and marine invertebrates are isomolal to seawater
(they are at osmotic equilibrium with seawater)
Body fluid concentrations in marine teleosts and lampreys are
300-350 mmol kg-1 water flows outward from the blood and
tissue fluids (hyposmolal)
Cartilaginous fishes and coelacanths retain urea and other
nitrogen-containing compounds, raising osmolality of their blood
slightly above that of seawater (hyperosmolal)
Exchange of Water and Ions

What is the
osmolality of
freshwater verts?
 Exchange of Water and Ions

Salt ions (e.g., sodium, chloride) can also diffuse through the
surface membranes of an animal (mostly gills)
Water and salt balance of a marine vertebrate is constantly
threatened by outflow of water and inflow of salt (ions)
In freshwater, vertebrates are threatened by inflow of water and
outflow of salt
Most fishes are stenohaline can tolerate only modest changes in
salinity
Some fishes are euryhaline and can move between freshwater and
seawater (water and salt gradients are reversed as they move from
one medium to the other)
 Freshwater Vertebrates Teleosts and Amphibians

Although body surface of fishes has low permeability to water and

ions, the gill surface has high permeability most water and ion
movements take place across gill surfaces
Water is gained by osmosis and ions are lost by diffusion
To compensate for water influx, kidneys of freshwater fishes or
amphibians produce large volumes of urine
Salts are actively resorbed to reduce salt loss
Large glomeruli produce copious urine, but ultrafiltrate is isomolal to
the blood and contains essential blood salts
To conserve salt, ions are reabsorbed across the proximal and
distal convoluted tubules by active transport
Urine becomes hyposmolal water is removed from the body
and ions are conserved
Salts from food compensate for ion loss, and teleosts have
ionocytes in the gills that actively take up chloride ions (sodium
ions also enter the gills, passively following Cl-)
 Freshwater Vertebrates Teleosts and Amphibians

Amphibians face similar osmotic problems as freshwater fishes

Entire body surface is involved in active uptake of ions from the
water via specialized cells (lose ions by diffusion)
Acidity inhibits active transport of ions in both amphibians and
fishes, often causing death in acidified habitats
 Marine Vertebrates

Osmotic and ionic gradients of marine vertebrates are essentially

the reverse of those in freshwater vertebrates
Seawater is more concentrated than the body fluids, so there is a
net outflow of water by osmosis and inward diffusion of ions
As in freshwater, marine teleosts experience most exchange across
the gills
Kidney glomeruli are small, and glomerular filtration rate is low
(little urine is formed)
Urine is more concentrated but remains hyposmolal to the blood
To compensate for osmotic dehydration marine teleosts drink
seawater
Sodium and chloride ions are actively absorbed across the lining
of the gut, and water flows by osmosis into the blood
To compensate for salt load, chloride cells in gills pump chloride
ions outward against a large concentration gradient
 Marine Vertebrates

Cartilaginous fishes maintain internal concentration of body fluid

close to that of seawater
Retain nitrogen-containing compounds (e.g., urea) to produce
osmolalities slightly hyperosmolal to seawater (gain water by
osmotic diffusion across the gills)
Gills of cartilaginous fishes have low ion permeabilities
Do not have well-developed salt-excreting cells in gills, but
secrete ions from rectal gland
About 5% of cartilaginous fishes are euryhaline, and one genus of
sharks (Glyphis) and stingrays of the family Potamotrygonidae are
exclusively freshwater

Rare Bornean river shark

Kidney structure
and function in
freshwater and
marine teleots
Water and salt regulation
by freshwater and marine
teleosts
 Nitrogen Excretion by Aquatic Vertebrates

Proteins and amino acids contain nitrogen, which is reduced to

ammonia during metabolism through deamination
Ammonia is soluble in water and diffuses readily, but is extremely
toxic
Most vertebrates excrete nitrogen as a mixture of nitrogenous waste
products:
Ammonotely ammonia: (bony fishes, excrete ammonia
through skin, gills, urine), produced by deamination and requires
no metabolic energy
Ureotely urea: (mammals are mostly ureotelic), urea synthesis
requires more energy than ammonia production, but urea is less
toxic than ammonia, and can be concentrated into urine
(conserves water)
Uricotely uric acid: (reptiles, including birds are mostly
uricotelic), pathway for uric acid is complex and requires more
energy than synthesis of urea most effective way to excrete
nitrogenous wastes and conserve water
 Responses to Temperature

Temperature affects rates at which chemical reactions proceed

Q10 = the ratio of the rate of a reaction at one temperature and the
rate of the reaction at 10C higher
Q10 of 1 means rate is constant
Q10 of > 1 means rate increases
Q10 of < 1 means rate decreases
Standard Metabolic Rate (SMR) of an organism is the minimum
rate of oxygen consumption needed to sustain life
SMR is temperature sensitive energy cost of living is affected by
changes in body temperature
Maintaining temperature differences in aquatic habitats requires
well-developed thermoregulatory mechanisms
Responses to Temperature

Swimming speed of a goldfish

 Responses to Temperature

Activity of an enzyme from

a lungfish

Spontaneous activity of goldfish

 Responses to Temperature

Poikilothermy body temperature is variable

Homeothermy body temperature is stable
Traditionally, fishes, amphibians, and reptiles were considered
poikilothermic, but terms are not useful for describing groups of
organisms
E.g., many fishes live in water that fluctuates less than 2C, and
some mammals and birds allow their temperatures to drop
considerably during certain times of the year, or daily
Ectothermy gain heat largely from external sources (e.g.,
basking)
Endothermy gain heat from metabolic production to raise body
temperatures
Also, not mutually exclusive birds and mammals are primarily
endothermal, but some species make use of external sources of
heat (e.g., roadrunners, contra brooding pythons)
Note: metabolic production of heat is costly
 Responses to Temperature

Regional Heterothermy refers to different temperatures in

different parts of the animals body
Several fishes maintain parts of their bodies at temps 15C
warmer than surrounding water, a remarkable feat considering
that each time the blood passes through the gills, it comes into
temperature equilibrium with the water
Some sharks: countercurrent blood flow using retia mirabilia,
where heat from warm venous blood leaves muscle tissue and
lies adjacent to countercurrent, cool arterial blood from the gills
Scombroid fishes: arrangement of retia that retains heat
produced by myoglobin-rich swimming muscles near vertebral
column
Billfishes: warm only the brain and eyes using a specialized
muscle that only produces heat (does not contract) superior
rectus eye muscle modified with many mitochondria
Responses to Temperature
Responses to Temperature