The critical thermal maximum of theiguanid lizard Urosaurus ornatus
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Authors Tremor, John William, 1932-
Publisher The University of Arizona.
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THE. CRITICAL THERMAL MAXIMUM OE THE IGUAN3D
LIZARD UROSAURUS ORNATUS
by .
John W® T rem or
A Thesis Submitted-, to the Faculty of the
DEFARTMEMT OF ZOOLOGY
In P a rtia l Fulfillm ent of the Requirements For .the Degree Of
; DOCTOR OF PHILOSOPHY
In the Graduate C ollege'
THE U W ERSITY OF ARIZONA
1962.
THE UNIVERSITY OF ARIZONA
GRADUATE COLLEGE
I hereby recommend that this dissertation prepared under mydirection by _____ John W» Tremor_______________________________entitled THE CRITICAL THERMAL MAXIMUM OF THE IGUANID LIZARD
UROSAURUS ORNATUS________________________________
be accepted as fulfilling the dissertation requirement of the degree of Doctor of Philosophy_____________________________
3c, /9& 2-Dissertation Director Date
After inspection of the dissertation, the following members of the Final Examination Committee concur in its approval and recommend its acceptance:*
GrK-'uli. 3 i m
l-Q .nn.G u.*
*This approval and acceptance is contingent on the candidate's adequate performance and defense of this dissertation at the final oral examination. The inclusion of this sheet bound into the library copy of the dissertation is evidence of satisfactory performance at the final examination.
STATEMENT BY AUTHOR
This thesis has been submitted in partia l fulfillment of r e quirements for an advanced degree at The University of Arizona and is deposited in The University Library to be made available to borrowers under ru les of the Library.
Brief quotations from this d issertation are allowable without special perm ission, provided that accurate acknowledgement of source is made. Requests for perm ission for extended quotation from or reproduction of this m anuscript in whole or in part may be granted by the head of the major department or the Dean of the Graduate College when in their judgement the proposed use of the m aterial is in the in terests of scholarship. In all other instances, however, perm ission must be obtained from the autnor.
SIGNED:
ABBtMAQT
The c ritica l therm al maximum of the tre e lizard U rosaum s
ornatns linearis (Baird and Girard) wag determ ined a f te r seasonal ■.
acclim atioa and therm al acclim ation in the laboratory. Laboratory
aoolim atioa eraslstocl of exposure fo r seven days # various constant
and ctiHraaily^Gyoleci t e n p o r a t a r e s * . ■
. f h ree responses to h e # s tr e s s w ere used to m easure the effeet
Of therm al acclimation,' /'These# in./the o rd e r o f th e ir occurrence^ a re •
(1) gape# (2) lo ss of rightiiig: response '(the c r itic a l the rm al maximum)#
and (3) death. Significant, correlations- were found between all th ree
th e rm al responses,, •
- - , Acclimation at li^C# 20*C# and 30*0 resu lts in increasing
c r itic a l therm al maxima (45,31^0# 46,. 39^0#. and 46*98*0 respectively).
Acclimation a t 36*0# n ear- eccritic temperature# seem s to be s tre ssfu l
since the OTM .falls to 46,86*0, The CTM values for diurnally-cycled .
anim als . (8*“'2T*0#. 16*-36*0# and, 29®«4©#C) a re significantly different
from corresponding"' values for consfant-tem perature acclim ation,
' In. th is iguanid# the hom w static function of behavioral therm o^
regulatiQn appears re la ted to c r itic a l th e rm al responses# inasmuch as
,th e re is a behavioral control of physiological compensation to tem perature
' i s . ' . . '
which, m asks the effect of seasonal tem perature change*
The effects' of ra te s of heating? and of repeated exposure to
high tem perature? on the c ritica l therm al responses w ere investigated
and a re briefly reported*
ACKNOWLEDGEMENT
The author is indebted to Dr. Charles H. Lowe, J r . whose
encouragement and advice concerning the problem he originally
suggested made possible this paper. The suggestions and help of
other professors and graduate students are also deeply appreciated.
Of particular benefit was the aid of Gerald O. Gates, Wallace G. Heath,
P ete r J. Lardner and Jam es R. Walker. The investigation was
conducted in the University of Arizona Arid Lands Program , supported
in part by a grant from the Rockefeller Foundation.
TABLE OF CONTENTS
U C q >e- vb .xct » , » a -P' i» - » 49 »• # j o » o o . & ^ -a ■*>'&■& a 1
& <b *> S> o o x> O 43 -e> 49 O O 4 0 4 > - ® « - © P - O p - O p « Q - O - O O Q P O O 0 j 9 4 ^ 4 > 0 0 0 » 0 p -O t i 2
REsiJLTfs a n d d i s c trssioN $ & <> <? -® ^.»»«©• .e P ^ ^ o ^ »* i s
]mTEEUkL.Fm IS
EXPERIMENTAL CONSTAWM^ACC 9 (> A & -a -® -® O ■& J0 -O Q o O « -O -O rO -0 .0 6= O fi> » P -O -0 -® p -© -0 -C3 -© -0 29
E X P E R m E N m L CYCLED-TEM PERATm ' '^ ^ .C z -a -o o -o a -e -p ^ o -o ho -o o p -®. -o & p p -o -® -o P p P o 9 -e 0 0 -® o o « -o 3 9
RELATION OF ACCLIMATION AND RESPONSE 43
THERMAL STRESS &-p. * * * -0 9-0 © * p -© & © © ^ © © © © © © © ©©©©©. © © © 56
ACCLIMATION AND WEIGHT CHANGE ©©© .©©p©©©©© 62
> P P -o # p. p. -f © -#. © 4 -d 9 <> © -0 , p © 9 © © © p p. e © © 9-6
EFFECT OF BATE OF HEATING * 7 2
SUMMARY AND CONCLUSIONS ......... 75
LITERATURE CI7? ED # -q a» #«»»^»-o ^«o ct»»»<>»9 < #» o -» 80
Page
X, Season of year (1961) and tem peratu re CC) ofoccurrence of the rm al response points » . 4 , , , 1 4
. 24 Comparison of spring (1961) and fa ll (1961)thermal response poxnts * -e o c o ® » & & & v ® © <, © © © © © ©»«© © © 18
3, One-factor analysis of variance for seasonaltherm al r^esponse ©©©'©©k©©©©©©© ©© ©©©©©©©© ©©©©©© »© ©©©© ©©© 19
4© R egressions and correlations of therm al responsepoints and season (.19 61 ©©©©©©.©©©©©.©©©©;©©©©©©©© ©«© © © © © ©
5© Tem peratures (°C) of therm al response points,according to season^ after acclim ation a t 30°C ©..»,«©©©.©» 24
6© T em peratures (% ) of therm al r esponse points^according to season, a f te r acclim ation a t 35°C . ©»© © ©»© © © © 25
7© Constant-tem perature acclim ation and. tem p era tu re ''■(0C) of occurrence of therm al response points ©©©©©©,»,©©, 32
8© Cycled-tem perature acclim ation and tem perature(0C) of occurrence of therm al response points ..,»©©© © © © © © © 40
9© Values for reg ression and correlation of theacclim ation index resulting from controlled(constant and cycled) tem perature acclim ation ©©,©©.„©»»©, 46
10© Acclimation index and mean temperature data forthe seven days preceding field collections and' thermal response determinations ©©©©»»©©©©©»©»,»©©»©©©©, 48
11© Mean body tem peratures recorded upon collectionin sum m er (May, 1952) and fall (Sept©, and Oct©, 1952)© ,©, 51
12© Regression and correlation coefficients for CTM andfor death on gape response following varyingconditions of acclim ation © © © © © © © © © © ©©©©©©©©© © © ©. © © © © © © © © © 55
TABLES (continued)
13*. R egression and correlation coefficients fo r CTM (n - 200) and fo r death (n « 1Y2) on gape response fo r field, constant-tem perature and cycled-tem pera- tu re acclim ation data *»********** * $ * * * * * * *»* * * # p & *»* 3 7
14, Comparison of the regression coefficients for gape r esponse v ersus CTM and gape response v e r s u s death by the method of least squares (b) and Bartlett*s method (B) * * * <* * * -* * ,»-p * * * * -»#»* * * ***,-* * * # * * * * * * * *, * * * 59
IS* Tem perature of gape response following alternating subjections to tem perature increase .and re tu rn to*
■ acclim ation tem peratu re (1#^C) * * * , * * * , , , , , , , , 60
16* Percent weight difference (loss o r gain) after sevendays acclim ation at GonsWnW emperature *,* ** . 63
17* Analysis for CTM diff erences in m ales and fem alesfollowing therm al acclim ation * * * * * * $ * *.*, * »• # * § * * -p * * * * * 66
IB, Snout-vent length (mm))and sex fo r the data inTable 17 ****** * * :*■ * * * * * p. * * * * * * p * * ******** * * * * ® * * ****** 67
19 * Regression of CTM on snout-vent length andcorrelations fo r cum ulative' data' sum m arized in iTables 17 and IB * * * * * * * * .* *, * * ************ * * * * * * * * * * * #. * 6B
20* T herm al response points and ra te of tem perature ■ -in c rease (after 30oG acclimation) ********************* * * 73
w i i
FIGURES
FIGURE
1, Chamber a i r tem perature record of 8* - 27®C cycled-tem perature experiment „ , -O Q ■© s » P @ -4> » p -© P 9 P P O -
2, Apparatus used in determining c ritica l therm al responses. Left# w ater foatli with testing cham bers.Right, te letherm om eter with probes 6
3, Urosaurus ornatns (male) in heat s tre s s chamber# removed from w ater bath (see Fig* 2)s showing body tem perature probe leading out from under tail# andthe a ir tem perature and surface tem perature probes*«, * 8
4, Seasonal differences in gape response and CTM .> , „ , 15
5„ Seasonal diff erences in acclim ation index andtem perature of d e a t h @ *. . * o . 18
6» Regressions of gape response and CTM on tim e 'of year > • . . . . . . . p. . Q. -o * . q f p * p. . . #. p *: 2.0
7« Regression of acclim ation index and tem peratureof death onfxme of year p . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.1.
8. Seasonal differences M and CTM.when acclim ated fo r seven days to 30°C and 3 5&C . . . . . . ■■ 26
9. Seasonal differences, in acclim ation index and ‘ tem perature of death when'acclim ated for sevendays a t 30 and 3 5* _*C.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2.7
10.. Regression of c r itic a l therm al maximum - on gap e *response for field acclim ation . . . . . . . . . . . . . » . . . . . . . . . 30
.11. Regression, of tem perature of death;on/gape 'response for field acclimation . . . . . . . . . . . „>,....». * . . . . . 31
12. Gape response and CTM as influenced by constant- ;tem peratu re acclmxat^on-. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
. v l l i ;'
FIGURES (continued)
FIGURE
IK Acclimatioii index and death a s influenced by■ constant-tem perature acclimation » * ^ ^ »„ * * #«*
: 14-p Regression of c ritic a l therm al maximum on gape . ■response fo r constant-tem perature acclim ation , #»,,»»«
15, R egression o f tem peratu re of death on gape response ■ fo r constant-tem perature acclim ation
1% Gape response and CTM as influenced'by cycled’-tem perature acclxmatron «* * .* «* #»-* <? # # * #■ * # # # •# p
17, Acclimation index and tem perature of death as influenced by cycled-tem perature acclim atien
18, R egressionof CTM on gaperesponse for ■cycled-' • tem perature acclxniation »0», ® e. <? a p», © -©© -©# © © © .©©©©© © © © ©
10© Regressiou of tem peratu re of death on gape response • for- cycled-tem perature acclim ation ©. © ©$ # ©,, © © ©#
20© R egression of acclim ation index on constant-tem pera- tu re and cycled-tem perature acclimation© Means of environmental air- tem peratures and the re sp e c tiv e : ■ . acclimation index plotted ©»©»© © * # .*■• © © #» »© © © © © © # »■ * © © © ©
21© Morning body tem peratures recorded upon, capture ■ in Sabino Canyon in May, 1052 and Sept© -Oct© * 1952© ©»„
22© R egressions of CTM and tem perature of death on gape response by method of least squares and B artlett’s method including a ll field, constant- tem peratu re and cycled-tem perature acclim ation
FIGURES (continued)v
FIGURE
23, Succeeding gape responses with body tem perature increases alternating with decreases to the acclim ating tem perature of 15°C, (a) Increase to tem perature of preceding gape response, (b) Increaseto 2*C above preceding gape tem p era tu re ,, , , , , . * *«, *. *, * ■ 61
24, Percent weight lo ss a fte r 7 days acclimation a tdifferent acclim ation t empera tu res , , , , , , , * , , . , , , , , , * . , , * 64
25, Comparison of fem ales and m ales in relation to CTMand snout-vent length to r seasonal acclim ation, , 69
26, Comparison of fem ales and m ales in relation to CTM and snout-yent length for constant-tem perature acclim ation ,, ^ ^ , 70
27* Comparison of f em ales and m ales in relation to CTM and snout-vent length to r cycled-tem peratur e a c c l i m a t i o n , , 71
28, Response to heating after 30*0 acclim ation a t a tem perature increase of 0, 6^C/min, and of 0 ,2 C! / m m , , , , , , *, *, * , , , , , * * , , , , , , , , , , , , , , * * * 74
a
INTRODUCTION
Urosaurus ornatus (Baird and Girard) is a sm all iguanid tre e
lizard that is abundant in parts of Arizona# neighboring sta tes and in
northern Mexico. Many aspects of its general and physiological ecology
have been under investigation in the Southwest during recen t years . The
present study was undertaken to investigate c ritica l therm al phenomena
in a natu ra l population of th is species and to com pare the findings for
na tu ra l seasonal ra te levels with those fo r experim ental tem perature
controlled acclim ation.
It was established by Lowe and Vance (1955) that the c ritica l
therm al maximum of th is diurnal rep tile responded significantly to
experim ental therm al acclim ation. Thus the present work was designed
for (1) a determ ination and evaluation of the effect of natural seasonal
events on the c r itic a l therm al maximum of this species 5 and (2) fu rth er
clarification of the nature of the upper therm al response points as well
a s the natu ra l sequence of th e ir occurrence under experim ental con
ditions.
Since M osauerts (1936) early investigations of reptilian therm al
behavior and the m ore extensive and well-known work of Cowles and
Bogert (194% considerable attention has been given to the subject of
therm oregulation in rep tiles and in o ther coldblooded organism s.
Several investigators have pointed to the evolutionary implications of
therm oregulation in lizards (Bogert, 1949; N orris, 1953; Vance, 1953;
Pearson, 1954; Cowles, 1958; Stebbins, 1958; Inger, 1959; Larson,
1961) . Cyclic activity and therm oregulation in rep tiles was recently
reviewed by Saint Girons and Saint Girons (1956).
Recent review s of therm al acclim ation and compensation have
been given by Bullock (1955), F ry (1958), and P ro sse r (1958, 1961).
With the early work of F ry (1942), indicating the significance of tem pera
tu re acclimation in the physiology and ecology of fishes, m ore such
studies became concerned with coldblooded vertebrates in generaL Yet,
i t was but relatively recen t that experim ental tem perature acclimation
was brought to bear specifically on the problem of rep tilian therm al
relations (Vance, 1953; Lowe and Vance, 1955; Dawson and Bartholomew,
1958; Dawson, 1960).
METHODS
Specimens of the subspecies Urosaurus ornatus linearis Baird
w ere collected from a population occurring in lower Sabine Canyon,
Santa Catalina Mountains, Pim a County, Arizona. This locality is nine
airline m iles (13. 8 m iles, by road) from the laboratory a t the University
of Arizona, in Tucson. The canyon stream supports a riparian woodland
of ash, oak, sycam ore, cottonwood, m esquite and willow with adjacent
slope covering of climax desert vegetation dominated prim arily by
foothill paloverde, sahnaro, and associated plants of the Arizona Upland
section of the Sonoran D esert, The anim als w ere taken from 2800 to
3400 feet; and along the edge of the canyon stream w here desert and
riparian vegetation interm ingle.
A to ta l of 312 anim als was collected by the thread-noose method.
This number includes those used in prelim inary experim ents but not
reported on here; data for 253 individuals a re reduced and analyzed.
Total body weights of the anim als ranged from 0.47 to 5.20 gram s.
Both m ales and fem ales were represen ted in each experim ental sam ple.
Upon collection the lizards w ere brought to the laboratory,
weighed, and either immediately tested for therm al tolerance o r placed
under controlled environmental conditions. F o r the la tte r, the anim als
were identified by m arking with fingernail polish after weighing, and
were then placed in battery ja rs in lots of six o r le ss . Before the ja rs
were placed in the acclimation cham ber, they were supplied With w ater-
filled watch g lasses and a libera l number of mealworm la rvae (Tenebrlo
m olito r) which were constantly available to a ll experim ental anim als,
^Precision" Low Tem perature Cabinets (B, O.D, boxes), with
a tem perature control of plus or minus 0.3*0, and rela tive humidities
ranging from 45% to 60%, were used for m ost of the constant tem perature
acclim ation experim ents. F o r som e sam ples an Aminco-Aire Humidity-
Tem peratur e Control Apparatus (’’acclim atizer'r) was used,. This had
an aecuracy of 0« l'°C and rela tive humidity was se t a t 55%,
Experim entally-cycled tem perature acclimation,, with simu
lation of the therm al component of the natu ral environment* was accom
plished by th e u se of a walk-in incubator in which tem peratu re was
cycled front' the low experim ental extrem e to the high over a 2.4 hour
p e r io d .' The curve for 8* - 27°C (Fig, 1) is representative of tem perature
change in a ll such experimentsj the data fo r program m ing w ere derived
.from a ir tem perature recordings (Tempscribe* 7 days) a t the study a rea .
These tem peratu re cycles w ere experimentally controlled by a Partlow
program m er and w ere accurate to within 3 ,0°C .
The lizards were tem perature-acclim ated for a period of seven
days (except in one specified case) in a ll of the experiments using
controlled environments, A four watt fluorescent tube was placed within
a foot of the animal-containing jar* and light was supplied for the num ber
of daily hours corresponding to the daylight hours of the season of
collection.
A fter collection o r acclimation* the critica l the rm al maxima
and other therm al response points w ere determ ined by subj eeting the
anim als to heat stress* The experim ental apparatus (Fig, 2) consisted
Fig. 1. Chamber a ir tem perature record of 8° - 27°C cycled tem perature experiments
6
Fig. 2. Apparatus used in determining critical therm al responses. Left, water bath with testing cham bers. Right, teletherm om eter with probes.
of two battery ja rs j each lined with standard plant p ress cardboard,
im m ersed in a 1000 watt P recision Scientific Co* metabolic water bath.
The therm istor-sensing end of a sm all animal probe (Yellow Springs
teletherm om eter interchangeable probe^ No, 402) w as inserted into the
cloaca and past the pubis j the probe was fastened to the ta il by adhesive
tape, thereby allowing for freedom of body movement (Fig. 3). The
constant u se of this probe, m easuring directly the deep body tem perature,
elim inates experim ental e r ro r inherent in the procedure of assuming
the body tem perature to be that of the environment, o r in the procedure
of removing the anim al from the stressing environment to determ ine the
c r itic a l the rm al response point . One o r the other of these difficulties
has complicated sev era i previous studies and th e ir re su lts .
Also taped within the chamber was a Yellow Springs surface
tem perature probe (No* 409) to m easure substratum tem perature, and
an a ir tem perature probe (No. 405). A ll tem peratures w ere read from
m ulti-channel teletherm om eters (Y, S. model 46TUC) accurate to within
1% of range (11. 0#C), i. e . , ± 0. 055*C.
Heat s tre s s was imposed by allowing the body tem perature to
r is e , from that of the environmental tem perature, a t a ra te of approxi
m ately 0* 6°C per minute to the tem perature of death. This heat r ise
closely followed the a ir tem perature increase and fell somewhat below
the substratum ra te (0. 7°C /m in .). Although the ra te of tem perature
8
Fig. 3. Urosaurus ornatus (male) in heat s tre ss chamber, removed from water bath (see Fig. 2), showing body tem perature probe leading out from under tail, and the a ir tem perature and surface tem perature probes.
increase of both the a ir tem perature and the substratum tem perature
tend to fa ll off somewhat a t the experim ental extrem es (ca„ 48° - 50°C)?
body tem perature continues to r is e linearly and #onstantly» This pro
cedure of allowing the body tem perature to closely approxim ate the ra te
of tem perature increase of the environment has been followed since 1956
in this laboratory and since I960 in the present study* It is the experi
m ental condition which Hutchison (1961) also maintained for the determ i
nation of the c r itica l therm al maximum in salam anders; and as he sta tes,
the anim al is properly ’’heated, from a previous acclim ation tem perature
at a constant ra te just fa s t enough to allow deep body tem perature to
follow environmental te s t tem peratures without a significant tim e lag, ”
The tem peratures of occurrence of seven c ritica l therm al
response points w ere observed and recorded. A fter a certain increase
in the anim al's activity, as the tem perature begins to r is e from the
in itial environmental level, the f irs t notable response is that of gaping.
The animal suddenly opens wide its mouth and resp ira to ry movements
notably increase . The evaporative cooling function of this mechanism
seem s obvious but little definite inform ation supports the supposition
(Templeton, 1960). This response has the advantage of an objectively
and very accurately determ ined tem perature of occurrence. Hind leg
paralysis follows (ca. 3* 5°C higher), leaving the anim al capable of now
desperate movement only by its still-coordinated front legs. It is between
the points of hind leg and front leg paralysis that the loss of righting
response occurs- This point is determ ined by turning the animal on
its back and recording the tem perature a t which it can no longer righ t
itself- This is taken a s the c ritica l therm al maximum which herein
after will be re fe rre d to also as the CTM- Cowles® definition (Cowles
and Bogertj 1944) of the c r itic a l therm al minimum and maximum and
its la te r modification by Lowe and Vance (19 55) applies well to this
in terpr etation since ju st beyond th is point the anim al can no longer
"escape from conditions that will promptly lead to its death-,f More
closely conforming to this definition would be, perhaps, the therm al
response point following the loss of righting response, i- e , , front leg
paralysis. This point is , however, m ore difficult of m easurem ent and
is only recorded in 70% of the experim ents w hereas the loss of righting
response is recorded in virtually 100% of the cases-
The point occurring afte r front leg paralysis (ca, 0 ,2aC higher)
is a response of spasmodic movements, thrashings of the body and sharp
la te ra l movements of the head. This iS apparently the c r itica l therm al
response point that has been taken as th e CTM by m ost of the investigators
in this field (Cowles and Bogert, 1944; S tebbins,' 1954; Lowe and Vance,
1955; M cFarland, 1955; Rosenthal, 1957; Hutchison, 1961; Larson, 1961),
The anim al, when rem oved immediately a t th is point, will recover; its
chances of survival decrease the longer i t is left a t th is and higher
tem peratures* Although this reaction does not always take place a t the
tem peratu res expected* as has been noted also fo r salam anders (see
Hutchison* 1961)* if has been designated by others as the GTM not only
for its ecological significance but also fo r its ease of observation under
conditions w here the anim al could not be manipulated* and manipulation
is necessary to determ ine the loss of righting response and som etim es
front leg paralysis*
In th is species loss of righting response and onset of spasmodic
movements a r e points that bracket the fron t leg paralysis response*
'The fo rm er is considered the m ore appropriate for the GTM since* a t
onset of body spasms* fo r p ractica l purposes the anim al has already
lost its ability to escape from its lethal environment* Thus the GTM
here employed is the loss of righting response; th is occurs a fte r hind
leg paralysis and on the o rd er of 9 ,2°C lower than the f i r s t front leg
para ly sis .
About (X above initial spasmodic movements, strong whole-
body convulsions a re evident in about 50% of the cases. Death follows
within. 0„ 5o-l„,0oC and is determ ined consistently by the la s t movement
of throat o r eyelid. This is an a rb itra rily selected point that is*
nevertheless* a physiologically im portant one which became possible
to recognise a fte r the f i r s t few experim ents w ere conducted.
The c ritica l therm al r esponse points m ost consistently recorded
and given sta tis tica l consideration a re three; namely* gaping* less M
righting response (CTM)* and death. The strong correlation between
gape and CTM perm its calculation of an acclimation index* which is
the weighted mean of these two points* This combines the m ore
objective and consequently m ore p rec ise determ ination of the fo rm er
with the necessarily m ore subj ectively determined latter* The index
to some extent compensates fo r the natu ra l variability of the relation
between the responses* and possibly allows a m ore accurate com parison
between responses to differing conditions of acclimation* Although
Simply trea ted here as another th e rm al response* its ab strac t and
a rtif ic ia l natu re is c learly understood*
Statistical M e t h o d s Lowe and Vance (1955) modified Cowles1
and Bogertfs (1944) definition of the c r itica l therm al minimum or
maximum to the extent tha t it ,4may be visualized as a value tha t is
the arithm etic m ean of the collective the rm al points a t which locomotory
activity becomes disorganized and the anim al loses i ts ability to escape
from conditions that w ill promptly lead to its death*n Such means and
th e ir standard e rro rs have been determined* and they have been graphed
using th e method of S ic e and L eraas (1936)* Standard e r ro rs (±) follow
the values fo r means in. the Tables; the standard deviations a re not
tabulated* F o r determ ining values ©ft* the appropriate form ula given
in Simpson* Roe and Lewontin (1.960) was used*
Methods gf calculating reg ression and correlation coefficients^
and th e ir te s ts ? a re also taken from Simpson# Roe and Lewoniin (X960)#
and from others. Gape response is not an independent variable but is
determ ined by tem perature of acclim ation, B a rtle tt 's B (Bartlett#
1949) provides here a line of best fit and is used in determining the
relation between gape response# GTM# and death. F or purposes of
prediction# regressions by the method of least squares a re also calcu
lated,
F ra tio s determ ined from one-factor designs w ere also used in
ascertaining the effect of season on c r itica l therm al responses. Spe
cifically# te sts w ere made fo r the effect of the spring# summer# and
fa ll natural environmental conditions on the responses of gape# CTM#
and death# and on the acclim ation index.
F o r calculation and comparison# the means of the cycled
experim ental tem peratures and of the environmental field tem peratures
were used ,
RESULTS AND DISCUSSION
N atural F ield Acclimation
The tem peratures of therm al response points as influenced by
season a r e given in Table 1 and graphed in F igures 4 and 5, In a ll
Table 1
SEASON OF YEAR (1961) AND.TEMPERATURE (°C) OF OCCURRENCE OF THERMAL RESPONSE POINTS, DATA GRAPHED IN FIGS, | AND 5.
RESPONSE AND SEASON N MEAN RANGE
GapeSpring 23 42, 70 ± 0,15 41, 65 - 44, 60Summer 26 43,02 *■ 0,14 41,95 - 44, 70F a ll 10 43, 50 ± 0,28 41, 50 44, 50
CTMSpring 23 46,43 ± 0,10 45,30 - 47, 55Summer 26 46, 47 d: 0,11. 45,00 -4 7 ,5 5F a ll 10 46,35 ± 0e 16 45,20 - 47,20
Acclimation IndexSpring 23 44, 56 iWO, 10 43, 63 - 45, 60Summer 26 44, 75 A 0,10 43,60 - 45, 78F a ll 10 44,90 A 0,15 43, 85 - 45, 65
DeathSpring ' 13 48, 30 A 0,12 47,25 - 49,05Summer 26 48,37 a 0,12 47,15 - 49,80F a ll 10 48, 70 a 0,14 47, 60 - 49,30
GAPE
(°
C)
CT
M48 -
O 47 o
46 1
45
44
43
42
A A
41
SPRING SUMMER FALLT I M E ( S E A S O N , 1961 )
Fig. 4. Seasonal differences in gape response and CTM. Data in Table 1.
AC
CL
IMA
TIO
N
IND
EX
(°C
) D
EA
TH
5 0
Oo 49
48
47
46
45
4 444
43
SPRING SUMMERT I M E ( S E A S O N ,
FALL1961 )
Fig. 5. Seasonal differences in acclimation index and temperature of death. Data in Table 1.
17responses but CTM there is a trend over spring (Mar, - A p r,), sum m er
(June - July) and fa ll (Oct, ) ■ toward a higher therm al tolerance. This is
best shown by the tem peratures of gaping where the difference between
spring and fall is 0 ,80% which is significant a t the 1% level, It should -
be noted, however, that in no case does there appear a significant differ
ence in therm al response between spring and sum m er and between
sum m er and fall. Probabilities for the differences in the c ritic a l therm al
response points a re given in Table 2,
The th ree seasons studied (spring, sum m er, fall) w ere also
tested by analysis of variance, F ra tio s (Table 3) derived from one-
fac to r designs indicate a seasonal effect on gaping response ( P < 0 ol)
but show no such effect on the other responses. The lowest of these F
values is for the c ritica l therm al maximum (F 0.34),
When plotted on daily increm ents, and the reg ression lines
fitted, a trend in seasonal increase in therm al response levels is shown,
and again with the exception of CTM (Figs, 6 and 7), Regressions and
correlations a re calculated fo r a l l raw data, with n between 50 and 60;
the means of the c ritica l therm al responses, however, a re graphed for
a one to four day period. The probability values for the correlation
coeffecients a re given in Table 4,
The evidence cited above fo r an actual seasonal effect on
physiological adjustm ent is made m ore c lear by m easurem ent a fte r
18
Table 2
COMPARISON OF SPRING (1961) AND FALL (1961) THERMAL RESPONSE POINTS. DATA IN TABLE 1.
RESPONSE t P
Gape 3.5024 <0.01
CTM 0.4224 < 0 .7
Acclimation Index 1. 8074 < 0 .1
Death 2.1246 < 0 .0 5
19
Table 3
ONE-FACTOR ANALYSIS OF VARIANCE FOR SEASONAL THERMAL RESPONSES.
ESPONSE SOURCE SUM OF SQUARES D .F .
MEANSQUARE F P
ape Levels 4.46 2 2.23 3.84 c 0 . 10Deviations 32. 52 56 0.58
TM Levels 0.21 2 0.10 0.34 c 0 . 50Deviations 16.27 56 0.29
cclimation Levels 0.89 2 0.45 0.45 < 0 . 50idex Deviations 12.38 56 0.22
eath Levels 1.11 2 0.55 1. 77 CO. 50Deviations 14.50 46 0.31
UA
4 2
64 120- J 1----------- 1— 1—
MARCH APRIL JUNE JULYT I M E ( S E A S O N AND D A Y S :
190 1 rOCTOBER
Fig. 6. Regression of gape response and CTM on time of year. Data in Table 4.
AC
CL
IMA
TIO
N
IND
EX
(°C
) D
EA
TH
2150Oo
49
48 - o
47
46
45
44
4364 120 90
MARCH APRIL JUNE JULY OCTOBERT I M E ( S E A S O N AND D A Y S )
Fig. 7. Regression of acclimation index and temperature of death on time of year. Data in Table 4.
22
Table 4
REGRESSIONS AND CORRELATIONS OF THERMAL RESPONSE POINTS AND SEASON (1961). DATA GRAPHED IN FIGS. 6 AND 7.
IESPONSE b r P N
>ape 0.0031 0.306 <0.02 59
0.0003 0.043 < 0 .2 59
Acclimation Index 0.0007 0.156 < 0 .2 59
)eath 0.0025 0.230 < 0 .1 49
experim ental acclim ation a t a constant tem perature. The data a re given
in Tables 5 and 6. After acclimation a t both ;30QC and 35°C all of the
therm al response points w ere higher in the fall of 1961 than in the sum m er
of the sam e year (Eigs, 8 and 9). Again the la rger differences appear in ■
gape response. Here it m ust be stated that the 3 5°C acclim ated anim als
of the fa ll of 1961 w ere exposed accidentally to a rela tive humidity of
10%, The data^ although calculated here for purposes of comparison,
must be in terpreted with this in mind. Although data w ere gathered in
the fall (only) of I960, and tabulated and graphed, no com parative
seasonal data for that year is available.
Many w orkers have found m ore c lear-cu t seasonal differences
in some physiological responses of coldblooded verteb ra tes. Riddle
(1909) found ra te of digestion of a standardized object to change by about
30% in certa in months of the year in fish, frogs and tu rtle s . C arter
(1933) reported the tem perature-pulse ra te curve of the excised frog
heart to vary with the season, A photoperiodic, and thus seasonal,
influence on gonadal development was reported by Bartholomew (1953)
in Xantusia v ig ilis. Hoar (1955) found the goldfish to have m ore heat
resistance after exposure to a longer photoperiod. B essauer (1955)
showed a seasonal variation in size of liver* te stis and ovary in Anolls
carolinensis and Wilhoft (19 58) found the height of thyroid epithelium in
geeloporus oceidentalis to change over the seasons. Oxygen consumption
Table 5
TEMPERATURES (°G) OF THERMAE RESPONSE POINTS, ACCORDING TO SEASON, AFTER ACCLIMATION AT 30*C9
, DATA GRAPHED IN FIGS, 8 AND 9.
RESPONSE AND SEASON N MEAN RANGE
GapeF all, 1960 9 43.21 ±0*29 41. 70 - 44, 00Summer, 1961 9 43,25 ± 0,22 41.90 - 43, 85F all, 1961. 11 44.24 ± 0,19 43.00 45,40
CTM .F all, 1960 8 47.33 ± 0.13 46. 75 - 47, 90Summer, 1961 9 46,75 ± 0,12 46,00 - 47,20F all, 1961 10 46. 89 ± 0,15 46,00 - 47. 50
Acclimation IndexF a ll, 1960 8 45.39 ± 0,14 44.93 - 46.10Summer, 1961 9 45.00 ± 0.14 44,40 - 45,43
■ Fall, 1961 10 45, 58 ± 0 ,1 6 44, 60 - 46.25
Death .Fall, 1960 9 48.08 ± 0 ,1 8 47.00 - 48.80Summer, 1961 9 48.25 ± 0 ,1 6 47.10 » 49. 00F all, 1961 11 48,80 ± 0,17 47. 50 m 49,30
Table 6
TEMPERATURES (°G) OF THERMAL RESPONSE POINTS, ACCORDING TO SEASON, AFTER ACCLIMATION AT 3®^C,
DATA GRAPHED IN FIGS, 8 AND 9.
RESPONSE AND SEASON N MEAN RANGE
Gape ■ FaU, 1960 • 9 42*91 £ 0 ,2 4 41 .90 « 44, 50 .
Summer, 196:1 9 42,68 ± 0,39 41,35 - 45,25Fall, 1961 10 44*29 £ 0,37 42, 6 0 - 46.70
CTMF all, 1960 9 47, 58 £ 0,20 46. 75 - 48, 50Summer, 1961 9 46,14 £ 0,20 4 5 .2 0 -4 7 .3 0FaH, 1961 10 46,41 £ 0 .1 6 45. 70 - 47.20
Acclimation Index F an , 1960 8 4 5 .2 4 -fc 0,15 44, 65 - 46.25Summer, 1961 9 44,46 £ 0,29 43.28 - 46.27F a ll, 1961 10 45,39 £ 0.23 44. 65 — 46,30
Death , , . F all, I960 9 48,45 £ 0.21 4 7 ,3 0 - 49.00Summer, 1961 9 48*12 £ 0,20 . . 47,10 - 49,20F an , 1961 10 49, 46 £ 0,15 48,90 - 50, 60
GAPE
(°
C)
CTM
(°C
)
26
4 8
47
46
45
44
43
42
4 I J______ I__LFALL, I 960 SUMMER, 1961 FALL, 196
T I M E ( S E A S O N )
Fig. 8. Seasonal differences in gape response and CTM when acclimated for seven days to 30°C (open) and 35°C (solid). Data in Tables 5 and 6.
AC
CL
IMA
TIO
N
IND
EX
(°C
) D
EA
TH
27
5 0Oo
49
48
47
46
45
4 4
4 3FALL, I960 SUMMER, 1961
T I M E ( S E A S O N )FALL, 196
Fig. 9. Seasonal differences in acclimation index and temperature of death when acclimated for seven days at 30°C (open) and 35°C (solid). Data in Tables 5 and 6.
of the cunner was discovered by Haugaard and Irving (1943) to be higher,
when m easured a t 15^C, in the winter than in the sum m er„ At l^G, the
rev erse was found to be tru e fo r the salam ander Plethodon cinereus
(Vernberg, 1952),
In his extensive study of the c ritica l therm al maximum of
salam anders, Hutchison (1961) found the CTM of Diemictylus viridescens
to increase from spring to fall and decrease over the w in ter These
CTM were determ ined a fte r acclim ation a t 20°C fo r one month. These
findings a re in agreem ent with those of th is work, in showing what he
te rm s a lag in response to the higher sum m er tem peratu res. Such
re su lts may indicate a differential compensatory adjustm ent to the
acclimation tem perature over the seasons. This concept is discussed
in m ore detail below,
Larson (1961) found the fence lizard Sceloporus occidentalis
collected in the sum m er, in the C entral Valley of California, to have
a higher CTM than those collected a t the sam e locality in the spring,
Vance (1953), however, could dem onstrate no difference in oxygen
consumption in sum m er-collected U rcsaurus ornatus as compared with
anim als collected in the fall. This finding for resp ira to ry metabolism
in P rosaurus is the sam e as that found in th is study of c r itica l therm al
responses in the sam e species from the sam e locality .
Figures 10 and 11 show the regressions of CTM on gape
response and of death on gape response^ respectively* fo r a ll field data;
correlation coefficients a re given in Table 12, These correlations a re
Significant and indicate the consistency of the o rder in which the c ritica l
therm al r esponses occur and th e ir dependence on the tem perature of
acclim ation. This relatively sm all dependence of both CTM and tem p er-
a ture of death on acclimation tem peratu re for field animals# com pared
With the re su lts of experim ents involving constant-tem perature and cycled-
iem perature acclimation# is discussed below.
Experim ental C onstant- Tem perature Acclimation
Table 7 provides data# graphed in Figures 12 and 13# for the
tem peratures of occurrence of the c ritic a l therm al responses as
influenced by the experimentally imposed constant tem peratures of 15*0*
25*0 and 30°Ct The CTM (45, 31GC# 4 6 ,39°C# and 4 6 .98°C# respectively)
r is e s linearly.
Such dependency of CTM on acclim ation tem perature was
reported as early as 1895 by Davenport who worked with the tadpoles of
Bufo te r r e s t r i s , Hathaway (1927) was one of the ea rlie r workers who
investigated such tem perature to lerance a fte r acclim ation in fish.
Doudoroff (1942) found the heat resis tan ce of certain m arine fish to depend
CT
M
30
48
47 •o
46
45o SPRING • SUMMER A FALL
4 4454 442 43414 0
GAPE ( ° C )
Fi g. 10. Regression (solid line - method of least squares; broken line - Bartlettfs B) of critical thermal maximum on gape response for field acclimation. Data in Table 12.
DE
AT
H
31
50
49
•• o
48
47o SPRING • SUMMER a FALL
46
42 434 0 44 4541GAPE ( ° C )
Fig. 11. Regression (so1 id line - method of least squares; broken line - Bartlett’s B) of temperature of death on gape response for field acclimation. Data in Table 12.
T ab le 7
GONSTANT-TEMPERATURE ACCLIMATION AND TEMPERATURE f C) OF OCCURRENCE OF THERMAL. RESPONSE POINTS.
DATA GRAPHED IN FIGS. 12 AND IS;
END POINT AND ACCLBIATION TEMPERATURE N MEAN RANGE
Gape15% 19 42» 03 * 0. 23 40,55 - 44, 0525% 21 42.97 ± 0» 09 42 a 00 - 44a 0030% 29 4 3 .6 1 A 0.15 : 41a 70 - 45,4035% 18 42. 79 ± 0,22 41a 35 ** 45.25
GTM15% 19 4 5 .3 1 A 0.30 42.20 - 47.0025% 21 46-39 ± 0 .1 0 45.30 - 47.2030% 27 46.98 ± 0 .0 9 46.00 *< 47a 9035% 18 46.86 ± 0 .2 1 45.20 " 48, 50
Acclimatioii Index 15% 19 43. 67 ± 0.24 41. 52 - 45,3325% 21 44.68 ± 0P. 06 44,05 - 45.4030% 27 45.33 ± 0 ,1 0 44.28 - 46,1035% 18 44,85 ± 0 * 19 4 3 .2 8 - 46.27
Death .15% 14 47,49 ± 0 .2 3 44. 85 - 48,2025% 20 48.38 ± 0,07 47, 80 - 49,0030^C 29 48,41 ± 0.11 47.00 - 49.3035% T7 48.27 ± 0 ,1 5 4 7 ,1 0 - 49,20
48
47
Oo
46
O
45
44
43
Uo
LUCL<O
42
41
401_L
- I
J_J.
33
15 2 5 30 35A C C L I M A T I O N T E M P E R A T U R E - C O N S T A N T ( ° C )
Fig. 12. Gape response and CTM as influenced by constant-temperature acclimation. Data in Table 7.
4 9
OL 48
XH<UJ 47 Q
46
Oo 45
XUJo7 44
h-<
43
O 42 O <
4 I
■ i
34
- I
J_L15 25
A C C L I M A T I O N T E M P E R A T U R E30 35C O N S T A N T ( ° C
Fig. 13. Acclimation index and death as influenced by constant-temperature acclimation. Data in Table 7.
on acclim ation tem perature and Mellanby (1940) reported tem perature
acclimation-dependency of activity in Rana tem poraria and Salamandra
salam andra.;'
It was notj however, until the work by F ry (1942) and
co-w orkers on therm al acclim ation and its effect on fishes tha t the
effectiveness and significance of such acclimation became m ore gener
ally recognized. Since then a good deal of information on therm al
requirem ents and acclim ation in fishes, amphibians and rep tiles has
accumulated.
Over a se r ie s of acclim ation tem peratures^ and derived high
and low lethal tem peratures, B rett (1944) describes "to lerance polygons"
for certain freshw ater fishes within which the anim als survive, Doudoroff
(1942) found certain m arine fishes to acclim ate to a high tem perature
relatively quickly, losing the tolerance gained slowly a t lower tem pera
tu re s , I t may be noted h ere that the seven days laboratory acclimation
used in th is study of Urosaurus ornatus was chosen, not after a rig id
investigation of the tim e necessary for acclimation a t each tem perature,
but as a reasonable, convenient and unvarying length of tim e to be used
in a ll of the acclim ation experim ents. The general, desirability of a
much m ore detailed investigation of acclim ation therm operiods is, of
course, definitely indicated fo r m ost if not a ll anim al groups.
M cFarland (19 55) found therm al tolerances in Taricha tom sa
to be dependent on acclim ation temperature.. Both Zweifel (1955) and.
Hutchison (1961) listed a general relationship between the acclimation^
dependent CTM and the habitats of the salam anders investigated,
Lowe and Vance (1955) w ere the f irs t to show the effect of
therm al acclim ation on the CTM in a rep tile (U rosaurus ornatus),
Larson (1961) also dem onstrated th is effect in gceloporiis occidentalism
It is noted in Table 7 and in F igures 12 and 13 that the therm al response
points for U rosaurus a f te r 35°C acclim ation fa ll significantly below those
fo r 30°C acclim ation. It would seem that the constantly imposed n ea r-
eccritic tem perature of 35*0 (eccritic mean s 35, 5*0) is of a s tressfu l
nature. This conclusion is borne out by the work of Wilhoft (1958) who,
in Sceloporus occidentalism, ascertained the need of " re s t" periods from
prolonged exposure to "optimum" tem perature. When exposed for a
long period of tim e to th is tem perature, the height of the thyroid
epithelium increased^ and death eventually ensued. F reem an (1950)
found a drop in oxygen consumption of goldfish a t higher acclimation
tem peratures; the la tte r had a sim ila r effect on the CTM of certain
salam anders (Hutchison, 1961),
F igures 14 and 15 show the reg ressions of CTM on gape response
fo r a ll constant-tem perature acclim ation data. Here is shown, particu
la rly by the method of B artle tt5s B fo r regression , the g rea te r
49
48
47
O 46o
5
O 4 5
4 4
43
42
37
* 1 5 ° C• 25° C A 30° C a 35° C
4 0 41 42 4 3 4 4 45GAPE ( ° C )
Fig. 14. Regression (solid line - method of least squares; broken line - Bartlett's B) of critical thermal maximum on gape response for constant-temperature acclimation. Data in Table 12.
DE
AT
H
38
49 • A A## A ^ A ^
48
47
46
45
444543 4 44240 41
GAPE ( ° C )
Fig. 15. Regression (soUd 1ine - method of least squares; broken line - Bartlett’s B) of temperature of death on gape response for constant-temperature acclimation. Data in Table 12.
independence of the tem perature of death in relation to acclimation
temperature-, as compared with the CTM» This rela tively g rea ter
independence of the point of death is also illustrated in F igure 11. The
therm al point of death is apparently genetically determ ined in such a
way as to be non-adjustable.
Experim ental Cycled-Tem perature Acclimation
When the therm al environment of TJrosaurus is cycled diurnally
from 8* to 27QG3 16* to 36tiC? and 20* to 40*GS c ritic a l therm al responses
resu lt as given in Table 8 and in F igures 16 and 17. Insofar as is known*
this is the f irs t attem pt to determ ine therm al response points fo r a cold
blooded vertebra te a f te r acclimation to a tem perature-sim ulated natu ra l
environment. These tem perature cycles were program m ed according
to the natu ra l daily change from one extrem e to the other* and were
selected to represen t the tem peratures prevailing in the animals^ natural
environment during the spring* sum m er and fall* When therm al response
points as influenced by the cycled tem peratures (means) a re compared
with those resulting from corresponding constant-tem perature acclimation*
no differences a r e found (Tables 7 and. 8| F igs. 12, 13, 16, and 17).
Therefore* i t may be concluded that the means of these laboratory-cycled
.tem peratures conditioned the therm al response points.
Table 8
CYCLED-TEMPERATURE ACCLIMATION AND TEMPERATURE (*C) OF OCCURRENCE OF THERMAL RESPONSE POINTS.
DATA. GRAPHED IN FIGS. 16 AND 17,
................. r ■ ' '/END POINT AND ACCLIMATION TEMPERATURE - CYCLED (*G) N MEAN RANGE
Gape8* ~ 27°C 12 42, 50 ± 0P 22 41, 60 - 44, 50
16* * 36*C 22 42.8! 6 0,15 41,55 - 44.0520° - 40#C ■ 9 43, 37 * 0.12 4 2 ,80 - 43, 80
CTM.8° - 27°C 11 45,93 * 0,23 44,40 - 46,65
16* ~ 36*0 22 46,83 * 0,08 46,00 - 47, 4520* ~ 40*C 9 47,15 * 0,14 46.35 *- 47,65
Acclimation Index8* # 27 °C 11 44*23 * 0,19 43,00 45* 58
16* - 36*C 22 4 4 ,82 * 0,10 4 4 . 00 - 45* 7220* * -40*0 9. 4 5 ,2 6 * 0*09 44, 73 * 45* 60
Death8* - 27*C 9 4 7 ,5 7 * 0 ,1 9 4 6 . 40 - 4 8 . 10
'16* - 36*0 22 48* 24 * 0.11 47,10 — 48* 8020* r 4Q°C 7 48,47 * 0,15 48* 00 * 4 9 , 00
------ ---------.... ..... ........ ...— -------------- -
4 8
Oo
47
I -O
4 6
45
44
Oo
LUQ_<(£>
43
42
4 IJ—L
i
8 - 2 7 16-36 2 0 - 4 0(x = 17.5) ( x = 26) ( x= 30 )
A C C L I M A T I O N T E M P E R A T U R E - C Y C L E D ( °
Fig. 16. Gape response and CTM as influenced by cycled-temperature acclimation. Data in Table 8.
oo
4 9
Xh- 48 <LUQ
47
Oo
XLUO
46
45
I—< 44
OO 43
J__ L
I
1__L8 - 2 7 16-36 2 0 - 4 0
(x = 17.5) (x = 26) (x= 30 )A C C L I M A T I O N T E M P E R A T U R E - C Y C L E D ( °
Fig. 17. Acclimation index and temperature of death as influenced by cycled-temperature acclimation. Data in Table 8.
Figures: 18 and 19 (data in Table 12) show the regression of
CTM and tem perature of death* respectivelya on gape response fo r the
cycled-tem perature experim ents. The sam e conclusion may be drawn
here as that for the data graphed in F igures 14 and 15*
Relation of Acclimation and Response
Knowing the extrem es of a tem peratu re cycle (data presented
in preceding section) one can theoretically predict? from the regression
of c ritica l the rm al response on controlled acclim ation tem perature
(Table 9, Fig, 20), the various therm al ■response points; o r, knowing
one of the extrem es and the tem peratu re of a therm al response point,
the other extrem e can be predicted. This relationship may be expected
in, and is indeed shown by, a l l of the c r itic a l therm al responses but is
evidenced best by the acclim ation index, chosen here fo r the reasons
given above in the section on methods; see Table 10, Fig, 20 (in which
i t is plotted for a l l conditions of acclim ation). The data show, however,
th a t when m ean tem peratures fo r the seven days preceding each
collection in spring, sum m er and fa ll a re considered as influencing the
seasonal acclimation indices and these responses a re com pared with the
corresponding values fo r cycled-tem perature and constant-tem perature
acclimation, the tem peratures of these seasonal acclim ation indices (and
CT
M
a
49
48
O 47*o
45
4 4 -
43 4 4 45424 0 41GAPE ( ° C )
Fig. 18. Regression (solid line - method of least squares; broken line - Bartlett’s B) of CTM on gape response for cycled- temperature acclimation. Data in Table 12.
k5
o
Xh-<LUQ
49
48
o*47
• 16 - 3 6 ° C a 2 0 - 40°C
46
4 0 4241 43 4 4 45GAPE ( ° C )
Fig. 19. Regression (soUd Une - method of least squares; broken line - Bartlett’s B) of temperature of death on gape response for cycled-temperature acclimation. Data in Table 12.
T ab le 9
VALUES FOR REGRESSION AND CORRELATION OF THE ACCLIMATION INDEX RESULTING FROM CONTROLLED (CONSTANT AND C YCLED)
TEMPERATURE ACCLIMATION. DATA GRAPHED IN FIG.
s r P N
0.0982 0.9656 0,01 6
AC
CL
IMA
TIO
N
IND
EX
45.5Oo
45.0
44.5
44.0
43.5
FAl L A ^ no
C H - a S UMM E R
15 20 25 30 35ACCLI MATI ON T E M P E R A T U R E - C O N S T A N T
AND M E A N S OF E X T R E M E S ( 0 C)
Fig. 20. Regression of aeeHmation index on constant- temperature ( o ) and eye!ed-temperature ( o ) acclimation (35°C stress value plotted but not included in regression). Seasonal means of environmental air temperatures ( a ), calculated from low and high air temperature extremes, are plotted for the acclimation indices of the seasonally collected lizards. The squares (□ ) represent the same seasonal acclimation indices (y axis) plotted against seasonal body temperatures (the body temperature means plotted are for the mean difference between the diurnal eccritic body temperature and the nocturnal low body temperature extreme, which is taken as the low nocturnal air extreme); arrows indicate the same direction, in each case toward the regression line, for the plotted squares. Interpretation of the greater departure of summer values from the regression line is discussed in text.
fa b le 10
ACCLIMATION INDEX AND MEAN TEMPERATURE DATA FOR THE SEVEN DAYS PRECEDING FIELD COLLECTIONS AND THERMAL RESPONSE DETERMINATIONS* DATA GRAPHED IN FIG, 20,
FIELDt e m p e r a t u r e s SEASON TEMPERATURE (°C)
MAXIMUM MINIMUM m e a n ACCLIMATIONINDEX
Environm ental Spring 26,0 10,0 18,0 44.56(Air) Summer 40,0 23,0 31. 5 44, 75
F all 32,0 18,0 25,0 44,90
Body Spring . 35,5 10,0 23, 7 44,56Summer '36,8 23,0 29.9 44. 75F a ll 35,5 18.0 26 ,7 44.90
the other c r itic a l the rm al r esponses as well) a re hot a t a ll as predicted.
The points for seasonal acclim ation as determ ined by imm ediate environ
m ental therm al h istory fa ll considerably outside of the 9 5% confidence
lim its fo r reg ression of acclim ation index on cycled-tem perature and
constant-tem perature acclim ation.
This inability to predict a seasonal therm al response point from
knowledge of natu ra l environmental tem perature may not be surprising
in view of the now well-known mechanism of behavioral therm oregulation
(Cowles and Bogert, 1944). When one substitutes eccritic body tem pera
tu re m eans (as determ ined for U rosaurus by Vance, 1953) fo r the upper
environmental extrem es of spring, fa ll and sum m er, one obtains the
different values (see-.field body tem peratures; Table 10) as the mean
influencing tem peratu res. This calculated eccritic mean (34 .5*C fo r
spring and. fall, 3 6 ,8#C fo r summer) is the upper body tem perature
reached and controlled behaviorally by Urosaurus and, to an im portant
extent, is independent of the environmental tem perature. Assuming the
low extrem e body tem perature to be that of the environment (an a s
sumption made with little evidence) one finds the new m eans to bring the
values of the acclim ation indices fo r spring, sum m er and fa ll c loser to
the line of reg ressio n for controlled acclim ation (Fig, 20). Indeed, the
spring and fa ll values fa ll virtually on th is line. The sum m er value
lies s ti ll somewhat removed and may be a reflection of this very sm a ll
an im a ls lessened ability to control its body tem perature in the intense
sum m er heat above the ground surface. Its sm all size and consequent
rap id heat gain o r loss makes behavioral regulation m ore difficult and
eventually less effective. Vance's data (Table 11, Fig. 21) show this
difference in seasonal regulation. The possibility exists that the higher
tem perature extrem es of the sum m er may resu lt in a lower than expected
therm al response point, as evidenced by the lower response elicited by
35#C acclimation, plotted in F igure 20 for reference, but, of course,
not included in the controlled-acclim ation regression .
The proximity of a ll the seasonal body tem perature means
(behaviorally controlled over the seasons) and the proximity of a ll of
these experim entally derived c ritic a l therm al responses Is indicative
of the homeostatic function of behavioral therm oregulation in this species.
This is an im portant form of compensatory physiologic adjustment.
The subject of compensation and the limiting factors involved
continues to rece ive increasing attention. Much of th is work regards
compensation of ra te processes in poikilotherms to tem perature, as
recently reviewed by Bullock (1955). P rech t (1958) describes five m ajor
types of compensation ranging from none to complete; complete compen
sation being the maintenance of the sam e ra te function afte r a change in
the influencing factor. F ry (1958) points out that compensation, as a
rule, is only partia lly effective.
5 1
Table 11
MEAN BODY TEMPERATURE RECORDED UPON COLLECTION IN SUMMER (MAY, 1952) AND FALL (SEPT, AND OCT. * 1952)2 DATA FROM VANCE: (1953); DATA GRAPHED IN FIG, 21,
SEASON m e a n b o d y t e m p e r a t u r e s (°c ).(TIME, A .M .)
8 ^ 9 9 - 1 0 10 - 11 i i - i2N
Ssm m er : 35.0 " 3 0 .5 : 37,5 ■ ; 36,4
'Fall / 32,2 34,4 34,9 : 35,4
$2
3 8
37
36 UiQC3H 35
LaJCL
y jG—
34
>- 33QOCD
32
9o
10o
8 - 9
IIo
16o
9-10TIME
7o
13o
JL
10-11
14o
II-I2N
Fig. 21. Morning body temperatures recorded upon capture in Sabino Canyon in May, 1952 (open) and Sept.fOct.,1952 (solid). Numbers above points indicate sample size. Data from Vance, 1953.
Concerning such physiological compensation to tem perature in
the coMMooded vertebrate., there is relatively little known. Wells
(1935) was among the f irs t to rep o rt what we now regard as partia l
compensation. He reported mud suckers acclim ated a t 33°C and tested
a t 2QeC and 22°C to have an oxygen cbnsumption lower than those main
tained a t 11°C» Vance (1953) found that U rosaurus ornatus acclim ated
a t 35^0 had significantly lower ra te s of oxygen consumption a t 15*0
than those maintained a t 8*0, Dawson and Bartholomew (1956) sim ilarly
found that geeloporus occidentalis a t 16*0 had a higher ra te of oxygen
consumption when tested a t 33*0 than those acclim ated a t the la tter
tem peratu re, 3h contrast* Gelineo and Gelinee (1956) repo rt what
amounts to an alm ost complete metabolic compensation in two species
of Lacerta,
A behavioral control of physiological compensation to tem pera
tu re in coldblooded vertebrates has been dem onstrated in p referred -
tem perature work, Garside and T ail (1958) found rainbow trout* after
acclim ation a t 5°C? 10*C# 150C and 20*CS to pref er the tem peratures of
16*C* 15*G, 13*0* and 11*0* respectively, Wilhoft and Anderson (1960)
found gceloporus occidentalis to p re fe r a lower body tem perature (30,1*0)
a fte r acclim ation a t 35*0 than afte r 12*0 and 25*0 acclim ation (33, 7*0
and 33.2*0 respectively). The authors speculate that th is lower p re fe r-
endum accompanies o r is responsible fo r reduced oxygen consumption
(as reported for th is species by Dawson and Bartholomew, 1956) to
prevent "burning o u t.n
In th is study perhaps the higher tem perature values of therm al
response points obtained a fte r constant-tem peratnre acclimation in the
fa ll as com pared to sum m er (Tables 5 and 6; F igs. 8 and 9) rep resen t
a physiological compensation to the differing environmental tem peratures
experienced over these two seasons* On the other hand, the lower accli
mation index of sum m er anim als, m easured upon collection, compared
with fall (Table 5- Fig. 5) may be indicative of a behavioral compensatory
adjustment. In any case? Vance's (19S3) finding of no difference in
oxygen consumption of fa ll ITrosaurus as compared with sum m er, taken
into consideration with the acclim ation data of this study , points to the
compensatory natu re of rep tilian behavioral therm oregulation. If we
modify the following statem ent by Edwards (1943) to include the concept
of behavioral therm oregulation, then we have a situation in the sand crab
s im ila r to that in U rosaurus; "There a re indications that the effect of
tem perature may vary with season, and in ternal (behaviorally induced?)
changes in the anim al may thus offset the environmental changes in
tem perature in the direction of preserv ing stable m etabolism through the
seasons, ” Ther e is little question that Bullock (1955) is co rr ect in the
assum ption that the wide distribution of natural acclim ation and rela ted
ra te compensation point to th e ir la rge scale ro le in the ecology and
evolution of poikilotherm s.
55
Table 12
REGRESSION AND CORRELATION COEFFICIENTS FOR CTM AND FOR DEATH ON GAPE RESPONSE FOLLOWING VARYING CONDITIONS OF ACCLIMATION, DATA GRAPHED IN FIGS* 10y 11, 14, 15, 18 AND 19 „
ACCLIMATIONCONDITION
METHOD OF LEAST SQUARES BARTLETT’S
N b • . £ P B
F ield
CTMX
Gaping75 0,2795 0,2139 0,1 0.2151
DeathX
Gaping
Constant tem p,
CTM
59 0,2657 0, 8615 0,01 0,2171
XGaping
IS 0,6213 0.5844 0.001 0,5311
' Death ■X
Gaping
Cycled tem p.
78 0,6129 0,8578 0,001 0.2626
CTMX
Gaping42 0,41.50 0,4470 0,01 1.1730
DeathX
Gaping38 0,3529 0,4559 0.01 0.4693
Table 13 and Figure 22 sum up the regressions of CTM and
tem perature of death for field* constant-tenaperature and eycled-temper*-
a tu re acclim ation. A t te s t fo r the difference in the regressions of
CTM vs. gape (B r 0.4139) and death vs* gape (B = 0* 3101), calculated
by the method of Bartletfc^s B, is significant a t the 1% level (Table 14)*
This g rea te r dependence of CTM on acclim ation tem perature may be of
survival value, as compared with the relative tem perature independence
of death. The environmentally influenced CTM, then, increases di
rec tly with rising, potentially lethal tem perature.
T herm al S tress
\Table 15 and F igure 23 show the resu lts of experim ents designed
to determ ine effects of heat s tre s s . Animals were acclim ated a t 15°C
for four days in an attem pt to lower the tem perature of the gape response.
This was done to m ore readily allow detection of a r is e in the therm al
level of th is response, as influenced by ensuing experim ental conditions.
A fter initially determining the tem perature of the gape response* the
lizards were alternately heated (at the ra te of 0 .6*C per minute) to the
tem perature of the immediately preceding gape r esponse and cooled to
the 15*C acclim ating tem perature a t ca, 0 ,8*C per minute. There w ere
no significant differences in gape response after such treatm ent (Table 15,
Table 13
REGRESSION AND CORRELATION COEFFICIENTS FOR CTM (N=200) AND FOR DEATH #=172) ON GAPE RESPONSE® FOR FIELD, . CONSTANT-TEMPERATURE AND C YCLED-TEMPERATURE
ACCLIMATION DATA. DATA GRAPHED IN FIG« 22,
METHOD OF LEAST SQUARES BARTLETT'S
N b r P B
CTMX
Gape
Death
201 0,4625 0,4752 0,001 0,4139
X. Gape
176 '0,3267 - 0,4443 ! 0,001 0,3101
58
4 54 2 4 44 34 0 41
Fig; 22. Regressions of CTM and tem perature of death on gape response by method of least squares (solid lines) and B artlett’s method (broken lines), including all field, constant-tem perature and cycled-tem perature acclim ation data. Data in Table 13.
Table 1.4
COMPARISON OF THE REGRESSION COEFFICIENTS FOR. GAPE RESPONSE VERSUS CTM AND GAPE RESPONSE VERSUS DEATH BY THE METHOD OF LEAST SQUARES
<fo) AND BARTLETTfS METHOD (B)„DATA SUMMARIZED IN TABLE 13.
. REGRESSION COEFFICIENT t P
b 1.4925 < 0 .2
. B
.......... ■■■ ------------- ---------------------------------1--------:
3,1242 <0 ,01
T able 15
t e m p e r a t u r e o f g a p e r e s p o n s e f o l l o w in g a l t e r n a t in g SUBJECTIONS TOTEMPERATURE INCREASE AND RETURN TO ACCLIMATION TEMPERATURE (15*0),
PATA GRAPHED IN FIG, 23, ,
MAXIMUM BODY TEMPERATURE CG)
DETERMINATIONNO, 1 MEAN RANGE
Tem perature of preceding 1 (Initial) 8 42, 75 ± 0,39 41,35 r- 44.80gaping response 2 8 42,64 ± 0,41 41.30 - 45.05
3 8 42, 26 ± 0 , 32 41,05 * 43. 704 8 42,15 ± 0 ,3 8 40, 55 i- 43, 40
Tem perature of preceding 1 (Initial) 8 43,02 ± 0 ,3 4 41.45 - 44, 65gaping response plus 2*C 2 7 43,96 ± 0,49 42.00 - 45. 70
3 6 42. 76 ± 0 ,2 3 41, 70 - 43. 654 5 42.41 ± 0, 30 41.60 - 43. 50
o 44
43
42
41
4 0
LUCL<O
n m
1 2 3 4T E M P E R A T U R E I N C R E A S E S
(a.)
46
^ 4 5Oo 44
43 -LU
< 4 2O
41
4 0
T E M P E R A T U R E( b . )
3 4I N C R E A S E S
Fig. 23. Succeeding gape responses with body temperature increases alternating with decreases to the acclimating temperature of 15°C. (a) Increase to temperature of preceding gape response, (b) Increase to 2°C above preceding gape temperature. Data in Table 15.
Fig* 23a), although a downward trend is evident. In th is experim ent a l l
the anim als survived. Since U rosaurus is often active a t body tem pera
tu res close to these levels* these resu lts a r e not surprising .
If, however, the body tem perature each tim e is ra ised to a
point 2*G above the gape tem perature, L e* * to the level of the predicted
CTM* quite different resu lts a re obtained. F igure 23b shows an imm edi
a te r is e in response a fte r the in itial determination* and a m ore signifi
cant drop after the second heating period. Also* the anim als die a t a
ra te approaching the LDgg afte r four exposures to the tem perature of
the CTM (Table 15), The resu lts re c a ll Selye?s (1950* and elsewhere)
concept of the alarm reaction and sequential stages of what he term ed
the G eneral Adaptation Syndrome,
It would appear, then, that the tem perature at which gaping
usually occurs does not ac t as a significant ’’s tre s so r” whereas the
somewhat higher tem perature of the c ritica l therm al maximum definitely
does. These resu lts a re quite s im ila r to those for salam anders as
reported by Hutchison (1961), ,
Acclimation and Weight Change
Table 16 a n d Figure 2 4 provide data concerning weight gained
or lost during acclim ation a t the constant tem peratures indicated. As
63
Table 16
PER GEET WEIGHT DIFFEREMGE (LOSS OR GAIN) AFTER SEVEN PAYS ACCLIMATION AT CONSTANT TEMPERATURE.
BATA GRAPHED JN FIG. 2:4, '
ACCLIMATIONTEMPERATURE: E m e a n
f '. ' ' .
RAKGE
15% 13 ~ 9 15 d: 09 86 5a 43 -Mo 53
25% 21 4,0 8 5 db 1< 47 — 8P 68 — —IS *85
36% 31 8.0 84 i: lo 52 -Ho 11 -26.17
35% 19 ^ 1 4 .7 8 * 1 ,9 6 : — Op 82. — —2.8, 41
WE
IGH
T LO
SS
(%)
64
2 0
10 -
5 -
i
015 25 30 35
ACCLI MATI ON T E M P E R A T U R E - C O N S T A N T ( ° C )
Fig. 24. Percent weight loss after 7 days acclimation at different acclimation temperatures. Data in Table 16.
previously stated, anim als were provided with mealworms and water
during the acclim ation periods. In one experiment the anim als w ere fed
before being placed at the acclim ation tem perature of 15°C ; these were
excluded from this determ ination as was a sam ple of anim als acclim ated
to 35°C in a cabinet where re la tiv e humidity dropped to 10% during the
acclim ation period e
It is c lear that the anim als held a t 15PC lost m ore weight than
those acclim ated a t 25#C because of poor eating, or fasting^ a t the lower
tem perature. Otherwise; the increased loss of weight with increased
tem perature of acclim ation reflec ts the increased ra te of metabolism,
and, in th is case, i t is an alm ost linear relationship.
Sex and Body Size
Tables 17, IB and 19 and F igures 2 5, 26 and 27 show the
relationships between body size, sex and c ritica l therm al maximum.
A ll of the acclim ation experiments a re included in these determ inations,
except for the 20® - 4Q®C cycled tem perature group where only two
fem ales were represented in the sample^ In a ll cases but one (spring,
1961 acclimation) the fem ales a re found to have a higher CTM than the
males* But in every sam ple, the mean snout*-vent length of the fem ales
is shorter than that of the m ales, a situation reflective of the population
Table 17
ANALYSIS FOR CTM DIFFERENCES IN MALES AND FEMALES FOLLOWING. THERMAL ACCLIMATION*DATA GRAPHED IN FIGS, 25, 26 AND 27.
CRITICAL THERMAL MAXIMUM
ACCLIMATION CONDITION SEX N MEAN RANGE
FieldFall, ?60 &*61 fem ale
m ale9
1547* 03 ± 0 ,14 46*48 ± 0 .1 8
46.20 - 47,85 4 5 .2 0 " 47,80
Spring, r61 femalem ale
7IS
46.26 ± 0 .1 8 ' 46* 53 ± 0.12
45. 30 - 46. 60 45, 60 - 47, 55
Summer, f61 fem alem ale
' 13 46. 66 ± 0.18 . 46. 31 ± 0,10
45.00 - 47,55 45. 50 ~ 46. 90
Constant tem perature 15*G fem ale
m ale8
1146.07 ± 0. 23 44.76 ± 0 .4 2
45.30 - 47,00 42,20 — 46, 60
25^C femalem ale
813
46.40 ± 0 .1 2 46.38 ± 0,16
46,10 - 47.15 45,30 - 47.20
30eC fem alem ale
1512
47.25 ± 0,07 46, 65 ± 0 .1 2
46.70 - 47.85 46.00 - 47.20
35°C femalemale
1018
47.02 ± 0.17 46. 52 ± 0,23
46.10 - 48,10 4 5 ,2 0 - 47,95
Cycled tem perature 8* - 2 T C female
m ale5 ;6
46,19 ± 0 .3 2 45. 72 ± 0.30
45.00 - 46. 65 44.40 - 46, 50
16^ *- 36^0 fem alem ale
9 . 13
46. 94 ± 0. 10 46,76 ± 0. 12
46. 55 - 47. 40 46.00 - 47,45
67
Table 18
SNOUT-VENT LENGTH (mm) AND SEX FOE THE DATA IN TABLE 17.. DATA GRAFHED M FIGS, 25, 26 AND 27, •
SNOUT-VENT LENGTH
ACCLIMATION CONDITION BEX I MEANi
RANGE
F ie ldF all, * 60 & *61 fem ale 9 40, 22 ± 2*02 35 - 53
male 15 43,73 ± 1, 74 33 - 57Spring, *61 fem ale 7 41,14 ± 1, 98 34 - 48
male 15 46,07A 1,35 37 ~ 52Summer, *61 fem ale 13 49.31 ± 0 ,7 4 46 - 54
male 13 51.08 ± 0.67 4 8 - 56
Constant tem peratur e 15*C fem ale 8 44, 62 ± 2 .2 5 34 *• 50
m ale 11 47.82 ± 1.28 41 - 5325°C fem ale 8 48, 50 ± 1.07 42 - 51
m ale 13 51.23 ± 0 . 69 47 - 5530*C female 15 43. 67 ± 1. 79 34 — 52
m ale 12 45, 50 ± 2. 38 33 - 5535% fem ale 10 42.30 ± 2,06 35 - 51
m ale 18 44.39 ± 1 . 51 30 - 53
Cycled tem perature 8° - 27% fem ale 5 41.20 ± 2 , 50 3 5 - 4 7
m ale 6 48.83 ± 1 , 30 45 - 53
16* - 36% fem ale 9 47 .00± 1,02 43 - 51m ale 13 51,15 ± 0 ,8 6 45 - 55
68
Table 19
REGRESSIONS OF CTM ON SNOUT-VENT LENGTH AND CORRELATIONS FOR CUMULATIVE DATA SUMMARIZED IN TABLES IT AND 18V ■
SEX b E
t--- — -----‘ ’P'-"'
Fem ales -0,0229 0.2167 . ' 0.05
Males Oo0008 0.0059 ; 0.1
Fem ales and Males -0,0148 0.1179 0.1
CTM
(°
C)
S-V
LEN
GTH
(M
M.)
55
50
45
40
35
30
■A69
48
47
464
45
1 -
4 4
FALL, '60 8 '61 SPRING, '61 SUMMER,*6SEASON ( I 9 6 0 , 1961)
Fig. 25. Comparison of females (open) and males (solid) in relation to CTM (below) and snout-vent length (above) for seasonal acclimation. Data in Tables 17 and 18.
CTM
(°
C)
S-V
LEN
GTH
(M
M.)
55
50
45
40
35
30
- 470
- i
48
47
46
45
4 415 25 30 35
ACCLI MATI ON T E M P E R A T U R E - C O N S T A N T ( ° C )
Fig. 26. Comparison of females (open) and males (solid) in relation to CTM (below) and snout-vent length (above) for constant- temperature acclimation. Range for CTM of 15°C males extends to 42. 20°C. Data in Tables 17 and 18.
CTM
(°
C)
S-V
LEN
GTH
(M
M.)
55
50
45
40
35
30
■ . 71
48
47
46
45
448 - 2 7
A C C L I M A T I O N T E M P E R A T U R E16—36
CY CL E D ( ° C )
Fig. 27. Comparison of females (open) and males (solid) in relation to CTM (below) and snout-vent length (above) for cycled-temperature acclimation. Data in Tables 17 and 18.
72as a whole, Snout-vent length was taken as the p referred size m easure
ment ra th e r than weight since., in a ll the controlled acclim ation experi
ments ? weight changed variously during the period of acclim ation. The
correlation coefficient of length on weight fo r Urosaurus ornatus is
0. 991 ± 0.006 (Vance, 1953). This relationship is given by the equation
log W = log 0.001 - 3.290 log L.
That the differences in CTM evident here a re not due to sex but
ra ther to body size is suggested by the significant (negative) reg ression
of CTM on snout-vent length in fem ales (Table 19) . This finding alone,
however, may not completely ru le out the possibility of a true sex
difference.
In only two cases (Fig. 27) a re the differences in size between
the sexes significant. And in only two other cases (Fig. 26) a re there
sex differences in CTM, T herefore, it Is assum ed that combining the
data for the sexes, fbr quantitative analysis, is valid.
Eff ect of Rate of Heating
A fter acclimation a t 30°C, c ritica l therm al responses were
determ ined a t the usual ra te of tem perature increase (0 .6°C /m in.) and
compared with those elicited by rheostat-controlled heat increase a t a
slow er ra te of 0 .2°C per minute. The resu lts a re given in Table 20
T able 2 B
THERMAL RESPONSE POINTS AND RATE OF TEMPERATHRE INCREASE (AFTER 30*C ACCLIMATION)^
DATA GRAPHED IN FIG, 28.
RESPONSE AND RATE OF TEMPERATURE INCREASE E MEAN RANGE
Gape0, G C /min« 11 44.23 d: Oo 19 43.00 - 45.40O^B^C/min. 9 43. 74 ± 0 ,2 6 42. 90 - 46. 90
CTM ■0» @®C/min« 10 4.6* 89 * 0*15 46* 00 - 46.900 .20C /m in, 9 46.34 ± 0* 09 46*00- 46*90
Acclimation Index9»6#C/mibou 10 • 45.58 ± 0*16 44*60 - 46.25 "0»2#C/m im 9 45* 04 db 0P16 44* 50 - 45* 85
Death0 ,6*C/min» : IT 48* 80 ± 0*17 47* 50 - 49*30
9 48* 15 ± 0* 19 47. 00 - 49* 10
BODY
T
EM
PE
RA
TU
RE
5 0
4 9
Oo 4 8
4 7
4 6
4 5
4 4
4 3
74
E
GAPE NORIGHTING
DEATH ACCLIMATIONINDEX
T H E R M A L RESPONSE P O I N T S
Fig. 28. Response to heating after 30°C acclimation; at a temperature increase of 0. 6°C/min. (open) and of 0. 2°C/min. (solid). Data in Table 20.
75
and plotted in F igure 28, The fa s te r ra te of heating yields higher values
in each case. Although the Reverse might be expected (e. g . , see Fry*
1942; Hutchison? 1961) due to a longer tim e allowed fo r some acclim ation
a t the slower rate? the lower responses actually found might resu lt from
a decreased tolerance to the higher tem peratures when exposed to them
for a longer period of tim e. Moreover? this is reasonable when i t is
recalled that acclim ation to 35*C yields therm al response points lower
than those obtained by acclim ation at 30*0 (see Table 7), The continuing
need fo r increased standardization of techniques in th is field is obvious.
Summary and Conclusions
The c r itic a l therm al maximum of the tre e lizard TJrosaurus
ornatus linearis (Baird and Girard) was determined upon collection in
the field in southern Arizona during spring? sum m er and fall? and afte r
therm al acclim ation in the laboratory. Animals w ere acclim ated fo r
periods of seven days to the constant tem peratures of 15*0? 25*0? 30*0
and 35*0? and to the diurnally cycled tem peratures of 8* ~ 27*0?
16* - 36*0: and. 20* - 40*0, A to ta l of 312 animals w ere tested? 253
reported on here.
Seven acclimation-conditioned responses to increased tem pera
tu re w ere studied. T hree of these w ere eventually used to evaluate the
effects of field and experimentally imposed environmental conditions;
in the o rder of the ir natu ral occurrence, these a re (1) gape, (2) loss
of righting response (the c ritica l therm al maximum)j and (3) death. An
acclim ation index, the weighted mean of the f irs t two responses above,
has also been found to be useful as a m easure of acclim ation.
Information concerning the effect on the c ritica l therm al
maximum (CTM) of ra te s of heating and of repeated exposure to high
tem perature was also obtained* as was com parative weight loss data
over the period of acclim ation.
The homeostatic function of behavioral therm oregulation in
Urosaurus ornatus was found to be m easureable by c ritica l therm al
response. Data a re presented to show that such therm oregulation m asks
the effect of seasonal tem perature change. J r fall and spring, c ritica l
therm al responses as influenced by m ean field diurnal body tem peratures
a re found to closely approximate those experimentally controlled. T her
moregulation, for reasons discussed, appears to operate m ore efficiently
in these seasons than in the sum m er.
Seasonal differences after acclim ation a t 30°C and 35*C a re
shown and the ir possible compensatory nature is speculated upon.
Animals acclim ated a t 15*C, 25*0 and 30*0 respond in a linear
m anner in CTM, yielding values Of 45.31*0, 46* 39*0 and 46.98*0
respectively. At 35*0 acclim ation the CTM fa lls to 46. 86*0, significantly
77below the value fo r 30°C» Thus the n ear-eco ritic body tem perature of
35^0, when imposed over a number of days, appears to be stressfuL
When anim als a re dium ally cycled for seven days over the
ranges of 8° - 27*0, 16* - 36*0 and 20* ~ 40*0, the c r itic a l therm al
responses a re found to differ insignificantly from the corresponding
values of constant-tem perature acclim ation. This relationship, in regard
to field tem perature re su lts , is discussed.
Significant correlations between the in itial c ritica l therm al
point and CTM and death wer e found under a ll conditions of acclim ation.
When the tem peratures of the c ritica l therm al maxima and of
death a re trea ted as variables plotted upon the tem perature of gaping,
no significant difference is found between these two reg ressions. When,
however, a ll of these therm al response points a re plotted on tem perature
of acclim ation, the regression (by the method of B artleW s B) of death
on gape is significantly lower than that of CTM on gape. The apparent
acelim atability of the lower, ecologically im portant c r itic a l therm al
points (gape, loss of righting response) and the rela tive tem perature
independence of the physiologically significant death point a re discussed.
As m easured by the successive tem peratures of occurrence of
gaping, a fte r repeated body tem perature increase to tha t point, tem pera
tu re a t gape (after 15*C acclimation) does not appear to unduly s tre s s
the anim al. Repeated exposure to the tem peratures of the critica l
therm al maximum* however* does appear to elicit an "a larm " reaction.
Under sim ila r conditions of rela tive humidity and feeding*
anim als acclim ating a t 35^C lost m ore weight than anim als acclimating
a t 30°G and the la tte r lost m ore weight than those a t 25#C, On the other
hand* 15*C acclim ating Urosaurus lost m ore weight than those a t 25*0
since* a t the lower tem perature* they presum ably ate le ss (or nothing
a t all) .
Mo correlation of body size and CTM was found, in m ales, A
Significant negative regression of CTM on Size in fem ales was found.
This provides some evidence that the higher CTM for f emales* as com
pared with those of males* reflects the consistently sm alle r sizes of
the fem ales collected fo r each experiments
C ritica l therm al maxima were determ ined fo r 30*0 acclim ated
anim als by raising the body'tem perature 0,2*0 p e r minute. This was
compared to the determ ination made over a 0,6*0 per minute ra te of
heating (the standard ra te used throughout th is study) of identically
acclim ated lizards. The CTM of the sam ple m ore slowly heated a t 0.2*0
per minute was 46,34*0 which is significantly lower than the 46,89*0
fo r the sample heated a t the faste r ra te . The rev e rse might be expected
on the basis of e a rlie r assum ptions fo r fishes and amphibians; i. e. *
longer heating a t a slower ra te might yield a higher CTM. However* the
g reater amount of tim e spent a t the higher and m ore s tressfu l tem pera-
tu res (which do yield lower CTM; see Table 7, Fig. 12) apparently
offsets any acclim ation resulting from slow er heating.
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