The Respiratory system

(Kardong)
Respiratory Organs

Gills
-designed for water breathing
-they are dense capillary beds in the
branchial region that serve external
respiration
-supported by branchial arches
o Internal gills
-associated with pharyngeal
slits and pouches
-ventilation mechanism:
involves muscular pump of
the buccal cavity actively
driving water across the
internal gills
-often covered and protected
by:
Interbranchial
septum
-soft skinfolds
-chondrichthyan
fishes
Operculum
-firm
-osteichthyan fishes
o External gills
-arise in the branchial region
as filamentous capillary beds
that protrude into the
surrounding water
-found in larvae of many
vertebrates, including
lungfishes, some
actinopterygians, and
amphibians
-ventilation mechanism:
specialized muscles sweep
the gills back and forth
Lungs
-designed for air breathing
-elastic bags within the body
-embryologically arise as
endodermal outpocketings from the
gut
-connected to the outside by trachea
(branches into bronchi which

branches into bronchioles); entrance

is glottis guarded by tiny sets of
muscles that open and close it
-exhalation process forces most of
the spent air from lungs, some
remain in the passageways and mix
with the fresh air that enters
o Dead space
-the volume of used air within
the respiratory passageways
o Tidal wave volume
-the total volume inhaled in a
single breath
Gas bladders
-retains pneumatic duct, its
tubular connection to the
digestive tract
-filled with air that enters via the
pneumatic duct or with gas
secreted into the bladder from
the blood
o Swim bladders
-used to control the
buoyancy of the fish in the
vertical column
o Respiratory gas bladders
-heavily vascularized to
participate in supplementary
respiration
o Actinopterygian fishes
-a single elongated sac located
dorsal to the digestive tract
-difference between lungs and gas
bladders:
o Gas bladders are usually
suited dorsal to the digestive
tract whereas lungs are
ventral
o Gas bladders are single,
whereas lungs are usually
paired

-similarity of lungs and gas bladders: both

are outpocketings from the gut or pharynx
and have roughly equivalent nerve and
muscle supplies

The Respiratory system

(Kardong)
-agnathans, elasmobranches, or
placoderms have neither lungs nor gall
bladders

Cutaneous respiratory organs

-respiration through skin
-can take place in air, in water, or in
both
-common in amphibians
Breathing and embryos
-among amniotes, respiration
generally takes place directly
between the surrounding
environment and the embryo across
the skin
o Chorioallantois
-the membrane beneath the
hell of birds and reptiles that
acts as the respiratory organ
-supplies the respiratory
needs of the chicken embryo
for most of its time and the
egg

Ventilatory Mechanisms

Cilia
-line the routes along which the
water current flows
-their coordinated sweeps drive
water through the pharynx and
across the gills
-found in small aquatic organisms
with low metabolic demands, such
as protochordates
-remain in large vertebrates to clear
surface debris that can foul the
breathing device
o Surfactant
-reduces surface tension at
the water-air interface;
surface tension can collapse
the resulting microscopic
compartments in which gas
exchange occurs

-lowers this surface tension,

helps stabilize these
compartments, and
maintains their structural
integrity as elaborated
surfaces for respiratory
exchange
Muscular mechanisms
o Amphibians
-muscles within or associated
with the bases of the
projecting gills contract to
wave the gills back and forth
through the water
o Ram ventilation
-a technique by which a fishs
own forward locomotion
contributes to gill ventilation
o Muscular pumps
-actively drive water or air
through the respiratory organ
Water ventilation
-dual pump
-buccal and opercular
pumps
Air ventilation:
pulse pump
-fishes and
amphibians
Air ventilation:
aspiration pump
-involves rib cage and
muscular diaphragm
as pumps
-found in terrestrial
amniotes reptiles,
mammals, and birds

Phylogeny

Agnathans
o Larva of lamprey
-depends on cilia-lined
channels to gather food
collected

The Respiratory system

(Kardong)

-feeding ventilation current of

water is produced by pumps
composed of muscular velar
folds or velum and by
compression and expansion
of the branchial apparatus
-there are usually seven
pairs of slits; flaps of skin
cover these openings, which
act as valves
o Each gill includes a central
partition, the interbranchial
septum, that supports a set
of primary lamellae (gill
filaments) which is divided
into secondary lamellae that
contain respiratory capillary
beds; uses countercounter
system
o Adult lamprey
-water exits and enters the
pharyngeal slits
-muscle compression and
relaxation of the branchial
apparatus drive water in and
out
o Hagfishes
-scrolling and unscrolling of
the velum, one on each side,
together with synchronized
contractions and relaxations
of the branchial pouches,
produces a current of water
that enters via nostrils and
nasopharyngeal ducts,
flows in one direction across
the gills, and exits through
the common branchial duct
Elasmobranches
-gills lie lateral to the branchial arch;
each gill consists of a central
portion, the interbranchial septum,
covered on each face by primary
lamellae divided into secondary
lamellae

-can be holobranch or hemibranch

-facing plates of lamellae on
adjacent gills constitute a
respiratory unit
o Sharks
-dual pump mechanism that
creates alternating negative
(suction) and positive
pressures to draw water in
and then drive it across the
gill curtain
-parabranchial
compartments reveal
efficiency of this dual pump
-dual pump mechanism of
the shark maintains a nearly
constant pressure difference
between buccal and
parabranchial compartments
-unidirectional irrigation of
gills
-countercurrent system
Spiracle
-embryologically the
first gill slit
-small opening
-carries a much
reduced hemibranch
-also known as
spiracular
pseudobranch
-may play a role in
chemical sampling of
the passing stream of
water
Bony Fishes
-cartilaginous operculum
-protective cover
-part of dual pump used to ventilate
the gills
-countercurrent system
-fishes that ventilate a gas bladder
do so by gulping and forcing fresh
air through the pneumatic duct
Overview of Fish Respiration
o Gills
Lamprey

The Respiratory system

(Kardong)

-compression and
expansion of the
branchial apparatus
Hagfishes
-scrolling of velum
Active fishes
-ram ventilation
Gnathostomes
-dual pump
-branchial arches and
their associated
muscles are the
central components
Lungs and Gas Bladders
-air-filled sacs arise early in
fish evolution and serve
respiratory and hydrostatic
functions
-respiratory function
predominates in lungfishes
and tetrapods
Lungfishes
-trachea arises from
the floor of the
esophagus and joins
the lungs in their
dorsal position within
the body cavity, a
location that is more
favorable to buoyancy
control
-lungs are subdivided
into faveoli
Actinopterygian
fishes
-hydrostatic function
-air-filled gas
bladders gives
buoyancy to the fish
body and help resist
its tendency to sink
Primitive teleosts
-gas bladders retain
connection with
digestive tract via
pneumatic duct

Advanced teleosts
-lost connection with
the digestive tract
-closed bags of gases
called
physocleistous
swim bladder
-have an oval, a
pocket at one end in
which gas is
absorbed
Gas glands
-from which gas from
the blood is released
into the bladder
-blood vessels form a
countercurrent
capillary arrangement
rete mirabile
Secondary function
of swim bladders
-sound detection
-sound producer or
resonator
Elasmobranches
-No swim bladders
-avoids sinking by
having cartilaginous
skeleton, pectoral
fins, heterocercal
tail, and oil
(squalene) consisting
of lipids and
hydrocarbons

Amphibians
-in modern amphibians, skin is the
major circulatory organ, and in some
species, it is the exclusive
respiratory organ
-moist skin
-thin keratin layer
-rich supply of capillaries within the
integument
o modern amphibians
-ventilation depends not on
ribs but on pumping

The Respiratory system

(Kardong)
o

movements of the throat to

irrigate gills or fill lungs
aquatic amphibians
-pharyngeal slits often persist
with internal gills
-feathery external gills are
also present
most have lungs for
breathing air
septal surface
-partitions form and
subdivide to increase
the surface area
exposed to incoming
air
faveoli
-compartments
divided by
interconnected septa
-differ from the alveoli
of mammals in that
they are not found at
the end of a highly
branched tracheal
system; faveoli are
internal subdivision of
the lung wall thatopen
to a common central
chamber
amphibian larvae
salamander larvae
-have both internal
and external gills
-pumping action of
throat irrigates
internal gills with a
unidirectional stream
of water across their
surfaces
-feathery external gills
are held out in the
passing current
anuran larvae
-employ buccal and
pharyngeal force
pumps to produce a

unidirectional flow of
water across the gills
and generate a foodbearing current
-basic mechanism of
amphibian gill
ventilation includes a
buccal cavity and a
pharyngeal cavity
separated from each
other by a valve, the
velum
-buccal cavity is
separated from the
mouth by oral valve
and from the nares by
an internal narial
valve
amphibian adults
-gills are lost
-cutaneous respiration
-lungs ventilated by buccal
pump
four stages of anuran
respiration:
1. buccal cavity
expands to draw
fresh air in
through the open
nares
2. glottis opens
rapidly, releasing
spent air from the
elastic lungs; air
streams across
buccal cavity with
little mixing and is
vented through
the open nares
3. the nares close,
and the floor of
buccal cavity
rises, forcing the
fresh air into the
lungs

The Respiratory system

(Kardong)
4. the glottis closes,
retaining the air
that had just filled
the lungs, and the
nares open again
-anuran respiration
(buccal cavity pump)
also play a role in
vocalization
*Similarity of anurans (amphibian) and
lungfishes (fish) in breathing is that they
push fresh air into lungs against pressure in
contrast with birds, reptiles, and mammals

Reptiles
-pharyngeal furrows and
pharyngeal slits appear during the
early embryonic development, but
they never become functional after
birth
-some groups require supplemental
cutaneous respiration, but for most,
paired lungs is meet their metabolic
demands
o Snakes and lizards
-include a single central air
chamber into which faveoli
open
-faveoli are reduced in the
posterior of lungs making it a
nonexchange region
o Monitor lizards, turtles, and
crocodiles
-a single central chamber is
subdivided into numerous
internal chambers that
receive air from the trachea;
internal chambers are
ventilated by respiratory
movement, whereas the
exchange of gas between the
faveoli and these chambers
appears to occur by diffusion

-filing of lungs is based on aspiration pump

mechanism
-presence of ribs and intercostal muscles
-occasionally, exhalation is passive
-glottis closed between breaths to prevent
premature escape of air
o Snakes
-lungs have a posterior
respiratory organ (faveoli)
and a posterior saccular
organ (avascular)
o Caimans and other
crocodiles
-diaphragmatic muscles and
abdominal muscles
o Turtles
-lungs and other viscera
reside in a single fixed cavity,
so any change in volume
alters the lungs
Mammals
-ventilation by aspiration pump
Changes in the shape of ribcage and
action of diaphragm contribute to
pumping mechanism
o Diaphragm
-crural, costal, and sternal
parts, all of which converge
on a central tendon
-lies anterior to the liver
o Ventilation
-bidirectional and involves rib
cage and diaphragm
o Gas exchange
-the respiratory passageway
repeatedly divides, producing
smaller and smaller branches
until they terminate in blindended compartments, the
alveoli, in which gas
exchange occurs
Birds
-cutaneous respiration is
insignificant
-two lungs ventilated by aspiration
pumps

The Respiratory system

(Kardong)
-no alveoli
-passages branch repeatedly and
eventually form numerous tiny, oneway passages, the parabronchi, that
permit air to flow through the lungs
-air capillaries
-nine avascular air sacs located in
bones and viscera are connected to
the lungs
-trachea is divided into
mesobronchi which do not enter
the lung but extend posteriorly to

reach the posterior air sacs; give rise

to latero, venrto, and dorso bronchi
as well as secondary bronchi and
finally leads to parabronchi
-no valves
-anterior and posterior air sacs