Nandana

Aquatic respiration of
vertebrates and
physiological implications
of aquatic respiration

By: Nandana Nayana Kumara

University of Kelaniya
Sri Lanka
Student # BS/A002/066
 Aquatic vertebrates rely on following
surfaces for gas exchange,
 Cutaneous body surface
 External filamentous gills
 Internal lamellar gills

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 Cutaneous respiration

 Ex- eels, some catfish, bullheads, some

fish larvae, Salamanders (ex-siren), frogs

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 Significance of cutaneous
respiration is determined by,
 Favorable ratio of body mass
to surface area
(folding of skin-ex hellbender
salamander, Telmatobius,
cylindrical ones has less
surface/volume)
 Thickness of the skin
 Proximity of capillaries to the
surface
 Whether the circulatory system
is well developed or not
 Body movements- prevent
boundary layer of low DO

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 Using external filamentous gills,
 Present in few vertebrates, usually during
periods of increased activity and O2 demand.

E.g.-
 Elasmobranch embryos have filamentous gills
that extend from their internal gill chamber into
the surrounding albuminous fluid.
 Adult male lung fish has filamentous gills on their
pelvic fins at the time of nest building.
 Many aquatic salamanders

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 Using external……cont’d

 Some cat fish, sturgeons, paddlefish and

climbing perch have filaments of vascular
epithelium in their branchial chambers for gas
exchanging when they air breath.
 Male African ‘Hairy frog’ develops numerous
highly vascularized filaments of skin during the
breeding season.
 Larval amphibians (tadpoles) have external gills,
(in some they disappear when they
metamorphose and in others it’s not)
 Adult Necturus (mud puppy)

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 Using internal lamellar gills,
 Most fish rely on internal lamellar gills for
respiration

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Cyclostomes (lamprey and hag fish)
 They have 6-14
pairs of gill
pouches that
extend from the
pharynx.

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 Elasmobranch and Teleost fish
 They have a
considerably
more elaborate
gill structure than
do cyclostomes.

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 Thin epithelial
cell layers of
lamellae are
separated by
pillar cells, which
have extensions
that contact
adjacent pillar
cells and
surround the
capillaries.

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 Active fish tend
to have more gill
filaments and
more secondary
lamellae than do
sluggish fish and
air breathing fish.

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 Gill surface area is
highly correlated
with body mass.
 Slope is usually
about 0.7-0.8
 Intercept varies for
different taxa

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 Diffusion path length
 The diffusion path length between water and
lamellar blood is also significant in determining
the rate of oxygen exchange, since influx is
inversely proportional to diffusion path length.
 As a result, path length is quite low in both active
fish and sluggish fish.
 Laminar boundary layer due to water flow over
the gill tissue and mucus layers also reduce the
gas exchange.

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 Types of gill ventilation
 Buccal-opercular
pumping
 In teleost fish, a
complex coordination of
expansion and
contraction of buccal
and opercular chambers
maintains an almost
continuous one-way
flow of water over the
gills.

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 Ram ventilation
 Elasmobranchs draw water into the buccal cavity by negative
pressure and then force the water through the gill filaments to the
outside. This type of forced ventilation is called as ram
ventilation.
 Also many active fish use ram ventilation as an alternative means of
gill ventilation. These active fish swim with their mouth open, and
water is forced over the gills without buccal or opercular pumping.
 The energy for water flow over the gills is derived from the swimming
muscles, rather than the buccal and opercular muscles.

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 Many fish utilize buccal opercular pumping
when stationary or swimming slowly and
use ram ventilation at higher swimming
speeds; there is a transition stage
between these two methods.

 Many active fish (e.g. Tuna) can’t maintain

sufficient gill ventilation by buccal-
opercular pumping and suffocate if forced
to stop swimming.

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 The rate of water flow over the gills is
related to the body size, the metabolic
demand, and the extent of O2 extraction
from the water.

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 Blood flow through the gill lamellae is
generally in a countercurrent direction
to the water flow because this provides a
high efficiency of O2 exchange from water
to blood, if the rates of gill water flow and
blood flow are suitably maintained.

 The optimal ventilation/perfusion ratio

is
about 10 to 20 because water has a lower
O2 carrying capacity than blood.

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 O2 extraction efficiency (E) of
gills
 Percentage of O2 removed from the
incurrent water flow, relative to the
incurrent O2 content
 E= 100(PiO2- PeO2) /PiO2
 Where, PiO2 is the incurrent PO2 and
PeO2 is the excurrent PO2
 Values for fish gills are typically 20-60%,
and a similar range is found for
invertebrates.

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 Gill water flow is determined by,
• resistance to water flow
• pressure gradient across gills
 The effect of gill resistance is much
greater due to high surface area of the
lamellae and narrow interlamellar spaces.
 So, gill water flow is primarily determined
by the gill resistance.

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 Anatomical dead space
 Not all the water flowing through the gills
is available for gas exchange.
 Some water flows out between the ends of
the gill lamellae form adjacent gill arches.
This fraction of gill water flow is called as
the anatomical dead space.

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 Physiological dead space
 Not all the water flowing between the lamellae is
necessarily involved in gas exchange.
 A wide spacing of lamellae, exceeding 2× the
water boundary layer thickness, allows the central
stream of water to pass through the gills without
exchanging any O2; this is called Physiological
dead space
 Another source of dead space is a nonoptimal
ventilation/perfusion ratio (VW/Vb).

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 Fick’s law of diffusion
 VO2=DA (C2-C1)/x
Where,
 VO2 is the oxygen consumption rate (mlO2/min)
 D is the diffusion coefficient (constant)
 A is the diffusion area
 (C2-C1)/x is the concentration gradient

 For a particular species A is increased only due to a

chronic increased demand of O2. For example the
lengths of external gill filaments are increased in some
amphibians in response to hypoxia.
 So the most appropriate way to increase oxygen uptake
at instant increased demand is increasing the
concentration gradient. Many fish increase this by
increasing both rate and stroke volume.
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 Regulation of respiration in
aquatic vertebrates
 Matching of respiratory ventilation with the metabolic
demand requires a regulatory system.
A respiratory system generally has,
• Pacemaker neurons – spontaneous activity, produce
basic respiratory cycle
• Respiratory regulatory systems with sensory detectors
and a motor effector system
 A sensory system monitoring the decline in blood or
branchial pO2 rather than pCO2 would be most
appropriate for aquatic animals because the pCO2
increase only slightly due to its high solubility.
 In fish pO2 receptors are usually in the buccal cavity,
gills, opercular cavity, arterial blood vessels, venous
vessels or brain.
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 Many stimuli other than pO2 and pCO2 influence
the respiratory rhythm, e.g., stretch of the
branchial cavity, osmotic stress, mechanical
stress, chemical stimuli, temperature etc

 Temperature has two important influences on

respiration.
 An increase in temperature lowers the oxygen
content of the water and at the same time
increases the metabolic rate because of higher
body temperature; both effects increase the
respiratory demand.

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 Physiological implications of
aquatic respiration

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 The physiological properties of water determine
many aspects of the design and functioning of
aquatic gas exchange systems.
 Water is dense and viscous, with a low O2
availability.
 The water flow over a gill is unidirectional to
avoid the energy cost of reversing the water
flow.
 This one way flow of water makes it
mechanically practical to have a countercurrent
flow of blood and this can greatly increase the
physiological efficiency of gas exchange.

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 The flow of water over the respiratory
surface should be about 20 times more
than the blood flow through the respiratory
surface.
 There is a high metabolic cost to aquatic
respiration because water is a dense,
viscous fluid.

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 The low O2 content of water (relative to
air) and high cost of ventilation generally
limits aquatic animals to having a low
metabolic rate.
 The high heat capacity of water generally
precludes thermoregulation by aquatic
animals because the metabolic heat is
rapidly dissipated into the water ventilated
through the gills.

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Thank you for your attention

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