ador, veiga - 2001 - activity metabolism in the lizard sceloporus occidentalis

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    ACTIVITY METABOLISM IN THE LIZARD

    SCELOPORUS OCCIDENTALIS

    A L B E R T F. B E N N E T T A N D T O D D T . G L EE SO N

    Schoo l of B iological Sciences, Un ivers ity of Ca liforn ia, Irv ine, Californ

    Accepted 12/17/75)

    Stan dard levels of oxygen consu mp tion an d oxygen consum ption an

    duction during and after burst activity were measured in the iguanid

    p us

    occidentalis. T h e activ ity cap acity of this animal is restricted; it s

    ous movem ent for only 1-2 min. Th e contribution of aerobic metabolis

    tivity is strongly therm ally dep end ent. Maximal levels of oxygen con

    achieved during activity at

    3 0 4 0

    C .

    At lower temperatures, significant

    oxygen up take , which app ear t o result from restricted ventilation. T

    aerobic increase above resting levels occurs a t 5C, preferred body tem p

    species. Repa ym ent of th e initial stages of oxygen d eb t is also mo st r

    Lactic acid concentration reaches high levels during activity, and its

    g r ea te st a t

    30 C.

    Anaerobic metabolism represents

    62 -82y0

    of th e e

    during bu rst activ ity, accounting for nearly all of th e carb oh yd rate ca

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    LBERT

    F ENNETT ND

    TODD T

    GLEESON

    accumulated lactate, and the capacity

    for further activity is strongly curtailed

    during the recovery period. These

    generalizations also pertain to th e ac tivity

    metabolism of many other reptilian

    groups, b u t not to all Bartholomew an d

    Tu cke r 1964; B en ne tt 1973b).

    Certain methodological difficulties

    hav e hampered th e examination of to ta l

    energy metabolism, both anaerobic and

    aerobic, during activity in the lower

    vertebrates. Anaerobic energy produc-

    tion can be estimated by the lactic acid

    concentration in whole-body homoge-

    nates, a procedure which avoids the

    during and after stim

    activity. These intervals

    to permit a n accurate

    con tributio n of aerobi

    In addition, they can y

    on th e therm al depende

    eratio n of oxygen con

    activity and of maxima

    Sceloporus occidenta

    fence lizard, is one of t

    reptiles in southern C ali

    in a gre at variety o

    animals are territorial a

    during the day on an

    site of o ld lumber or

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    CTIVITYMET BOLISM IN

    LIZ RD

    moths (Galleria sp.) and had access to

    water a d libitum. Th e animals remained

    healthy and active and were generally

    held in captivity for less than 1 wk.

    Animals were fasted for a t least 2 days

    before experimentation.

    In the experiments which determined

    oxygen consumption and lac tate produc-

    tion simultaneously during activity, a

    single animal was weighed and measured.

    Electrical leads were implanted

    in

    the

    base of its tail. T h e lizard w as then

    placed in a rectangular Lucite metabo-

    lism cham ber (volume 534 cm3)

    equipped with ports for air flow and

    decrem ent was used to d

    consumption according

    Depocas and H a rt (195

    The environmental

    opened and incurrent a

    po rts were closed, isola

    the airtight chamber.

    stimulated to maxim

    electrical shocks of lo

    livered with a Harvar

    the implanted leads.

    chamber was hit a nd s

    the animal, which respo

    of rapid running. Stim

    tinued for 5 min. Air s

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    68

    ALBERT F BENNETT

    AND

    TODD

    T

    GLEESON

    analyzer (see Depocas and Hart 1957).

    Oxygen consumption during each minute

    interval of activity and recovery was

    calculated according to the formula

    where

    V volume of chamber (cm3) volume

    of animal (cma)

    V, volume of gas sample removed from

    chamber (cm3)

    RH

    relative humidity in the cham-

    figure 1. The thermal d

    function is complex.

    consumption is very s

    ture dependent (Qlo

    and 35 C but is essent

    independent at 20-25 C

    Such zones of therma

    oxygen consumption

    viously observed in o

    Bennett and Dawson

    nounced decrement i

    consumption occurred

    ning: the average eveni

    below that in the af

    decrease is not signific

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    ACTIVITY METABOLISM

    IN

    A

    LIZ RD

    stimulation are reported in figure 2. The Qlo 30-35 C = 2.2, Q

    total amount of oxygen consumed during

    In addition to the gre

    the period of stimulation increases with

    consumption the rate

    increasing temperature up to

    35

    C :

    consumption increase

    Qlo 20-25 C

    =

    2.9 Qlo 25-30 C

    1.4

    levels is also temperatu

    ce lo ~o ru s occidental is

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    70

    LBERT F.

    BENNETT

    AND TODD T. GLEESON

    20

    C, maximal levels are not attained consumption are coinc

    until after the cessation of activity and activity burst at

    30-40

    stimulation. As body temperature in- acceleration of oxygen c

    creases, the rate a t which maximal rates ing the first minute of

    of oxygen consumption are achieved also great between 20 and 2

    increases until maximal levels of oxygen Ventilation during burs

    TABLE

    THE EROBIC I N C R E M E N T D U R I N G 2 M I N

    OF

    BURST ACTIVITY AND

    SCOPE (MAXIMAL

    MINUTE

    AEROBI C

    I N C R E M E N T )

    IN

    SCELOPO

    OCCIDENTALIS

    TIMULATED FOR MIN

    AEROBIC

    SCOPE

    AEROBIC

    I N C R E ~ N T

    Time

    TEW. URING ACTIVITY tion

    ( C) N

    (cma

    O d k hl) (cmr OdIg hl

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    ACTIVITY M ETABOLISM I N A LIZARD

    TABLE

    INCREMENTS ABOVE PRESTIMULATION

    LEVELS

    DURING MI

    STIMULATION AND

    MIN

    OF RE OVERY I N

    L ' S ~ ~ ~ ~CCIDENTALIS

    Inc r emen t Increment

    during during Change in

    Temp.

    St imu la t ion

    Recovery

    Inc re me nt R e c o v e 0

    ( C)

    ( c m J

    O r l k h l ) cma

    O z l k h l ) ( c m a

    O r l k h l )

    Nore.-Values shown are means of 5-8 animals.

    stricted at low body temperature: no

    costal movements are evident during

    Lactate formation

    the bout of bu rst a

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    7

    ALBERT

    P BENNETT

    AND TODD T. GLEESON

    blood most rapidly at 35

    C

    All of these

    iguanids have preferred thermal levels

    of 35-38 C The interrelationships of

    oxygen debt, lactate elimination, and

    recovery in reptiles have not been

    examined, however, and it would be

    premature to speculate about their

    interdependence or physiological signifi-

    cance.

    2 5

    The delay at low b

    in achieving maximal

    consumption is not

    pected, considering the

    logical systems involve

    port. A similar lag ha

    reported for the marin

    rhynchus crist tus Be

    The low oxygen con

    Sc eloporu s occidental s

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    ACTIVITY XETABOLISM IN A

    LIZ RD

    TABLE

    3

    WHOLE-BODY LACTATE CONCENTRATIONS

    OF

    SCELOPORUS

    OCCIDENTALIS BEFORE DUR-

    ING,

    AND

    AFTER SPECIFIED

    PERIODS OF

    AC-

    TIVITY

    AT

    35 C

    Lactate Concen tra t ion

    Condition IV h s s

    w t )

    Unstimulated. 4 0.40+0.04

    min active..

    4

    1.34+0.16

    2

    min active.

    4

    1.73 0.10

    5 min active. .

    4

    1.70_+0.25

    5 min active+

    5 min recovery. 6 1.82f 0.08

    NOTE.-Valueshown are means+SE.

    temperatures in active Scelopmus is not,

    examining aerobic wo

    aerobic increment duri

    The thermal depend

    activity metabolism in

    more complex than wa

    posed. A low temper

    over a wide range of b

    was postulated on the

    tions on whole-body

    (Bennett and Licht 19

    lactate concentrations

    Bennett et al. 1975). T

    of whole-body lactat

    narrow thermal increm

    and Dipsosaurus (Ben

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    c e l o ~ o r u soccidentalis

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    ACTIVITY METABOLISM I N LIZARD

    hydrate catabolized enters anaerobic

    pathw ays see Be nn ett e t al. 1975).

    Energetic ou tpu t during burst ac tivity

    is maximal over th e range of 30-35 C in

    S.

    occiderttalis. Body temperature of

    animals in the field appears to have

    some seasonal lability over this range

    McGinnis 1966). Groups of anim als

    caugh t in th e field during spring through

    fall have mean body temperatures of

    34.3-35.9 C. A similar group of winte r-

    active animals, however, had a mean

    body temperature of 30.4 C. All groups

    tested in the laboratory had preferred

    Sceloporus, in spite

    Dipsosaurus is 2-3

    Maximal anaerobic en

    almost identical in the

    production occurring a

    rus and 40 C in D

    output is maximized

    modes at preferred b

    40 C, in Dipsosaur

    output is consequent

    species 52 vs. 40 pmo

    this increment is as

    sup po rt of m uscular a

    locom otory efficiency

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    76

    ALBERT

    F BENNETT A N D TODD T

    GLEESON

    D A W S O N .

    R .

    1975. On the physiological signifi- cidentalis. Comp. Biochem

    cance of the preferred body temperatures of rep-

    FR Y F.

    E

    J 1947. Effects

    tiles. Pages 443-473

    i n

    D. M. GATES nd R.

    B

    animal activity. pu b. O

    SCHMERLds. Perspectives in biophysical ecol-

    8:1-62.

    ogy. Springer-Verlag New York.

    D A W S O NW.

    R.

    and G. A. BARTHOLOMEW.956.

    Relatio n of oxygen cons um ption to body weight

    temperature and temp erature acclimation in

    lizards

    Uta slansburiana

    and

    Sceloporus occi-

    dentalis. Physiol. 2001. 29:40-51.

    DEWCAS.F.. and T S HART.1957. Use of th e Pau l-

    M c G r N ~ r s N 1966. Scel

    ferred bod y temp era ture

    lizard. Science 152 : 1090

    IMOBERLY . R. 1968a. The

    the common iguana

    Igu

    under restraint. Comp.

    ing oxygen analy zer for measu remen t of oxygen

    consum ption of an imals in open-circuit system s

    19686. T h e metab ol

    and in a short-lag closed circuit apparatus

    mon iguana Igztana igzc

    J

    Appl. Physiol. 10:388-392.

    ing. Com p. Biochem. Ph

    FRANCIS . and G. R . BROOKS.970. Oxygen con-

    WILSON

    K. J

    1974. T h e

    sump tion rat e of heart beat and ventilatory

    supply for act ivi ty to bo

    rate of parietalectomized lizards

    Sceloporzrs oc-

    species of lizards. Copei