Respiration and Circulation in

Reptiles
Elizabeth Timpe
Herpetology - EEB 3265/5265
23 February 2010
Lecture Outline
Respiration and Circulation in
Reptiles
Oxygen transport
Diving in aquatic and semi-aquatic species
Adaptations for O
2
uptake
Pulmonary
Extra-pulmonary (or nonpulmonary)
Hibernation and respiration
Changes in blood flow through the heart
Characteristics of blood*
*will not be covered in todays lecture
Pulmonary Uptake of O
2
Compared to amphibians, O
2
uptake through the skin
in most reptile species is minimal
Few exceptions discussed in this lecture
Reptiles rely largely on lungs for gas exchange
Have large lung volumes
10x more volume compared to mammals of similar size
However, reptiles lungs are much simpler
lack aveoli
lung surface areas are only ~1% that of mammals of similar
size
Remember, reptiles have a much-reduced metabolic rate
1-10% that of mammals
Pulmonary Uptake of O
2
Most snakes have only one lung (the right one;
Fig 7-6, pg 276 in textbook)
Some primitive lineages have a smaller vestigial
left lung
Maina et al. (1998)
In the sandboa (Eyrx [Gongylophis] colubrinus)
Anterior half of the lung is used for gas exchange
Posterior part is an air storage organ.
Pulmonary Uptake of O
2
Adaptiveness of large lung volume in reptiles
Store large volumes of air in the lungs
Used for aerobic metabolism
Have periodic, irregular breathing
Allows for less frequent breathing
Saves a lot of energy
Do not have to continuously contract muscles to fill lungs
with air
Reduces evaporative water loss across the lung surface
Especially important in desert reptiles
Pulmonary Uptake of O
2
Periodic breathing pre-adapts reptiles
for diving
Even terrestrial or arboreal reptiles can
remain under water for long periods of time
Apnea during a dive in an extension of their
normal breathing pattern
Allows the marine iguana to readily adapt to its
diving lifestyle
Allows snakes, crocodilians, and turtles to stay
under water for long periods
Pulmonary Uptake of O
2
Some largely aquatic reptiles have evolved
additional adaptations for prolonged
submergence
Alligators
Submerged for up to 2 hours
Support aerobic metabolism from stored oxygen in
blood and lungs
Marine snakes
(Hydrophiinae and Acrochordidae)
Have unusually large lungs that stores large
amount of oxygen for long dives
Dont have to rely on anaerobic metabolism to
support activity
Pulmonary Uptake of O
2
Sea Turtles
Stay submerged the longest of all reptiles
Highest metabolic scopes of all reptiles
Can swim for long periods of time without
resting
Have complex lungs with large surface areas
and large volumes
*Read Lutcavage and Lutz (1997) review of sea turtle diving physiology
(see bibliography)
http://en.wikipedia.org/wiki/File:Hawksbill_Turtle.jpg
Pulmonary Uptake of O
2
Cheloniidae
(marine turtles)
Shallow water divers, seldom >300 m
Store O
2
in lungs, used during dive
Dermatochelyidae
(leatherback sea turtles)
Deep water divers, up to 1000 m
At great depths, lungs would collapse
Most O
2
is stored in the blood
Highest hematocrits
Highest hemoglobin concentrations
Highest myoglobin concentrations
Pulmonary Uptake of O
2
File snake (Acrochordus granulatus)
High blood volumes
High hematocrits
However high hematocrits = increases blood viscosity, decreases
blood flow rate, decreases O
2
flow to the tissues
Storing O
2
in the blood does not mean increased capacity for activity,
but may allow for increased time submerged
High O
2
affinity
High O
2
affinity = high tendency for hemoglobin to bind with O
2
Allows animal to take up more O
2
into the blood, but inhibits release
of O
2
from blood to the tissues
Lowest aerobic scopes of any snake
Therefore blood characteristics in the file snake are related
to prolonged dive time and not increased activity
Extra-pulmonary uptake of O
2
Some aquatic species of reptiles have surprisingly high
capacities for EP gas exchange
Sea snakes - O
2
uptake through skin
Aquatic turtles - O
2
uptake through the pharynx or the
cloaca
Bagatto and Henry (1999) sliders vs. softshell turtles
Sliders (Trachemys) - dependent on aerial breathing, little EP O
2
uptake, excrete CO
2
into water, have short dives (~5 min), take
multiple breaths when at the surface, high tolerance for lactic acid,
anaerobic metabolism if needed
Softshells (Apalone) - rely on EP O
2
uptake to stay active when
submerged, make long dives (12-23 mins), single breath of air when
surfacing, cannot tolerate lactic acid buildup, and therefore cannot
rely on anaerobic metabolism to stay submerged longer
Extra-pulmonary uptake of O
2
Australian chelid turtles have muscular
cloacal bursae
Sacs branching off the cloaca
Have muscles to pump water in-out
Huge number of papillae, increasing respiratory
surface area
Example: Rheodytes leukops
Can obtain all their O
2
underwater
Cloacal bursae surface area = 16x that of smooth surface of
similar volume
Dependent on O
2
levels in water (prefer colder, fast flowing
waters)
Extra-pulmonary uptake of O
2
The total amount of EP O
2
uptake is hard to
determine for most species
Estimates for inactive sea snakes = 5-22%
Boa constrictor = ~3%
Cutaneous uptake is more important to small
individuals or small species
High surface to volume ratio
Cutaneous uptake may increase up to 120% for
active snakes compared to inactive snakes
Extra-pulmonary uptake of O
2
Partitioned O
2
uptake among differ EP organs in turtles
King and Heatwole (1994) - Elseya latisternum
49% - buccopharyngeal cavity
33% - cloacal bursae
18% - through skin
Podocnemys
90% - cloacal bursae
Sternotherus
70% - through skin
30% - buccopharyngeal cavity
Bagatto et al. (1997) - Kinosternon and Staurotypus
Less than 10% of O
2
from water, 90% from air
Aquatic CO
2
exchange was much greater (~40%)
Hibernation and Respiration in Reptiles
For reptiles that hibernate underwater, cutaneous
gas exchange is the only process available
Costanza (1989) - garter snakes hibernate underwater
(up to 80 cm) in abandoned wells
Metabolic rates depressed by 80%
Took up enough O
2
cutaneously to remain aerobic
Did not accumulate lactic acid even though submerged up
to 5 months
If exposed to anoxic conditions, snakes died
Hibernation and Respiration in Reptiles
Freshwater turtles - some species can take
up significant amounts of O
2
cutaneously
Examples - softshell turtles, musk turtles, map turtles
Favor highly oxygenated water
Metabolize aerobically; low tolerance for lactic acid buildup
Freshwater turtles - some species hibernate in mud or
other anoxic/hypoxic conditions
Examples - painted turtles
Metabolize anaerobically; high tolerance for lactic acid buildup
Found much further north than softshells, musk, and map
turtles
Hibernation in anoxic environments is common
Geographic variation between populations of painted turtles in
their tolerance of anoxic conditions
Hibernation and Respiration in Reptiles
Tolerance of anoxic conditions in turtles is due to the
buffering capacity of their shells
Calcium carbonate released from shell neutralizes the lactic acid that
accumulates
Essential for surviving long winters underwater, buried in mud
Juvenile turtles dont have the buffering capacity of adults
Reese et al. (2004) - juv. snapping turtles, painted turtles and map
turtles had survival rates in anoxic water 1/3 that of adults of the same
species
Explains why turtles have a greater capacity to survive anoxic
conditions than underwater-hibernating frogs
http://upload.wikimedia.org/wikipedia/commons/9/90/Defensive_turtle.jpg
Circulatory Adaptations
Mammals have a completely separated
circulatory system, with no mixing of blood in the
pulmonary and systemic circuits
(veins--> r. auricle--> r. ventricle--> pulmonary artery-->
lungs--> pulmonary veins--> l. auricle--> l. ventricle-->
aorta--> systemic arteries)
Reptiles do not have the same type of separated
blood flow, with some mixing occurring
Questions
1) How is the pattern of reptilian circulation adapted to
the animals oxygen transport requirements?
2) Is the reptilian pattern less efficient than the
mammalian pattern?
Snake, lizard, and turtle
hearts are different than
those of crocodilians
Snake, lizard, turtle heart
(Figures, 3rd page of handout)
Two atria (auricles)
Single ventricle
3 chambers
Cavum venosum
Cavum arteriosum
Cavum pulmonare
Circulatory Adaptations
1) Ventricle relaxes, blood from veins -> right and left atria
2) Blood from right atrium -> cavum venosum; blood from
left atrium -> cavum arteriosum
3) Ventricle contracts, muscular ridge that separates cavum
venosum from cavum pulmonare is not pressed against
the wall of the heart, so blood flows over the ridge from
cavum venosum -> cavum pulmonare, blood ->
pulmonary artery -> lungs
4) Cavum venosum is empty, and oxygenated blood from
cavum arteriosum -> cavum venosum through
intraventricular canal. Muscular ridge is pressed
against the wall of the heart, completely separating cavum
venosum from cavum pulmonare
- There is no mixing of oxygenated and deoxygenated blood
Circulatory Adaptations
The reptilian heart is no less efficient than the
mammalian heart
Reptilian system allows for shunting of blood
into different pathways under special
circumstances
These shunts are of two kinds:
1) left to right shunt: results in recirculation of
more blood to the lungs
Important in aerial breathing
2) right to left shunt: results in redirection of
blood away from the lungs to the body, particularly
the brain
Important during apnea
Circulatory Adaptations
In crocodilians:
(Figures on last page of handout)
A completely divided ventricle with two chambers, superficially
similar to that of birds and mammals
The right aortic arch (to the brain) arises from the left
ventricle, while the left aortic arch (to the body) arises from the
right ventricle
Most of the deoxygenated blood in the right ventricle normally by-
passes the entrance to the left aortic arch, goes through the
pulmonary artery to the lungs. Left aortic arch receives
oxygenated blood from the right aortic arch through a connection
called the foramen of Panizzae
Exhibits right to left shunting during diving and left to right
shunting during surfacing
Circulatory Adaptations