effects of feeding on thermal selection...

35
Effects of feeding on thermal selection by Constrictor constrictor Item Type text; Thesis-Reproduction (electronic) Authors Combest, Wendell Lee, 1951- Publisher The University of Arizona. Rights Copyright © is held by the author. Digital access to this material is made possible by the University Libraries, University of Arizona. Further transmission, reproduction or presentation (such as public display or performance) of protected items is prohibited except with permission of the author. Download date 21/05/2018 15:19:58 Link to Item http://hdl.handle.net/10150/347972

Upload: vuanh

Post on 21-Mar-2018

223 views

Category:

Documents


1 download

TRANSCRIPT

Page 1: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

Effects of feeding on thermalselection by Constrictor constrictor

Item Type text; Thesis-Reproduction (electronic)

Authors Combest, Wendell Lee, 1951-

Publisher The University of Arizona.

Rights Copyright © is held by the author. Digital access to this materialis made possible by the University Libraries, University of Arizona.Further transmission, reproduction or presentation (such aspublic display or performance) of protected items is prohibitedexcept with permission of the author.

Download date 21/05/2018 15:19:58

Link to Item http://hdl.handle.net/10150/347972

Page 2: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

EFFECTS OF FEEDING ON THERMAL SELECTION

BY CONSTRICTOR CONSTRICTOR

by

W endell Lee C om best

A T hesis Subm itted to the F acu lty of the

DEPARTMENT OF CELL AND DEVELOPMENTAL BIOLOGY

In P a rtia l F u lfillm en t of th e R equirem ents For th e D egree of

MASTER OF SCIENCE WITH A MAJOR IN ZOOLOGY

In the G raduate C o llege

THE UNIVERSITY OF ARIZONA

1 9 7 6

Page 3: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

STATEMENT BY AUTHOR

This th e s is h as been subm itted in p a rtia l fu lfillm en t of r e ­q u irem en ts for an ad vanced d eg ree a t The U n iv e rsity of Arizona and is d ep o s ite d in the U n iv e rs ity L ibrary to be made a v a ila b le to borrow ers under ru le s of th e L ibrary .

Brief q u o ta tio n s from th is th e s is are a llo w ab le w ithout s p e c ia l p e rm iss io n , provided th a t acc u ra te acknow ledgm ent of sou rce is m ade. R eq u ests for perm ission for ex tended q u o ta tio n from or rep roduction of th is m anuscrip t in w hole or in part m ay be gran ted by th e head of the m ajor departm ent or th e D ean of th e G raduate C o llege w hen in h is ju d g ­m ent th e proposed u se of the m ateria l is in th e in te re s ts of s c h o la r­sh ip . In a l l o ther in s ta n c e s , h o w ev er, perm ission m ust be ob ta ined from th e au th o r.

SIGNED: .

APPROVAL BY THESIS DIRECTOR

This th e s is .h a s been approved on th e d a te shown below :

ROBERT B. CHIASSONP rofesso r of Zoology

Page 4: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

TABLE OF CONTENTS

LIST OF ILLUSTRATIONS

LIST OF TABLES . . . .

ABSTRACT . . . . . . .

INTRODUCTION . . . .

MATERIALS AND METHODS . . .

RESULTS ........................ . . . . . .

DISCUSSION AND CONCLUSIONS

REFERENCES............................... . .

Page 5: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

LIST OF ILLUSTRATIONS

Figure Page

-I* F loorp lan of g rad ien t cham ber w ith 12 u n its u sed/ a s p o s itio n m arkers (51cm x 51cm x 1 9 8 c m ) . . 5

2* S u b stra te tem p era tu res a t each p o s itio n in th e g rad ien t . . . 5

3 . S e lec ted stom ach tem p era tu res for boa #1 fed a t 1200 . . . '. 11

4 . S e lec ted stom ach p o s itio n s for boa #1 fed a t 1200 ................ ... - 11

5« S e le c te d stom ach tem p era tu res for boa #2 fed a t 1200 . . . . 12

6« S e lec ted stom ach p o s itio n s for boa #2 fed a t 1200 . . . . . . 12

7 . S e le c te d stom ach tem pera tu res for boa #3 fed a t 1200 . . . . 13

8 . S e le c te d stom ach p o s itio n s for boa #3 fed a t 1200 .......... 13

9 . . S elected , stom ach tem pera tu res for boa #1 fed a t 1700 . . . . 14

10* .S e lec te d stom ach p o s itio n s for boa #1 fed a t 1700 ..................... 14

11. ; S e lec ted stom ach tem pera tu res for boa #2 fed a t 1700 . . . . 15

12. S e le c te d stom ach p o s itio n s for boa #2 fed a t 1700 . . . . . . 1 5 '

13. S e lec ted stom ach tem p era tu res for boa #3 fed a t 1700 . . . .- 16

14 . S e le c ted stom ach p o s itio n s for boa #3 fed a t 1700 . . . . . . 16

15 . S e le c ted stom ach tem pera tu res for boa #1 fed a t 2200 . . . . 17

16 . S e lec ted stom ach p o s itio n s for boa #1 fed a t 2200 .................... 17

17 . . S e lec ted stom ach tem p era tu res for boa #2 fed a t 2200 . . . . 18

18 . S e lec ted stom ach p o s itio n s for boa #2 fed a t 2200 .................. 18

iv

Page 6: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

V

LIST OF ILLUSTRATIONS—C ontinued

Figure Page

19. ; S e lec ted s tom ach tem p era tu res for boa #3 fed a t 2200 . . . . . . 19

20 . S e lec ted stom ach p o s itio n s for boa #3 fed a t 2200 . . . . . . . 19

Page 7: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

LIST OF TABLES

Table Page

1 . M ean S e lec ted Stom ach Tem perature °C „ 9

2 . Voluntary Maximum T em peratures °C . . . . . . . . . . 9

Page 8: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

ABSTRACT

The e ffe c ts of feed in g upon g a s tr ic tem peratu re s e le c tio n of

sp ec im en s of C o n s tric to r c o n s tr ic to r w as in v e s tig a te d in a labo ra to ry

therm al g ra d ie n t. Fed sn ak es dem onstra ted a s ig n if ic a n tly h igher therm al- . -' i .

p reference than did nonfed a n im a ls . The sn ak es m ain ta ined a more or

l e s s reg u la r cy c le of tem peratu re s e le c tio n approxim ating 24 h o u rs .

E levated stom ach tem p era tu res u su a lly occurred in the la te even ing and

con tinued un til n ea rly m idnigh t, w ith th e lo w es t tem p era tu res recorded

in th e ea rly to la te m orning h o u rs . F eed ing did not a p p re c ia b ly a ffec t

th is b a s ic c y c le . All tem perature f lu c tu a tio n s were c o rre la ted w ith

stom ach lo ca tio n reco rd s and w ere due to the snake a c tiv e ly changing

h is p o sitio n in th e g rad ien t.

The vo lun tary therm al maximum w as found to be g rea te r in fed

sn ak es when com pared to nonfed s n a k e s . Time of fe e d in g , w hether

1200, 1700, or 2200, had l i t t le to no e f fe c t on e ith e r th e tem peratu re

se le c te d or the 24-hour c y c le s . ,

Page 9: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

INTRODUCTION

Ambient tem peratu re has been shown to have an im portan t

in flu en c e on the ra te of g a s tr ic d ig e s tio n in many a n im a ls . Tempera­

tu re e f fe c ts on feed ing and d ig e s tio n in f ish are w e ll kn o w n . Brett

an d H iggs (1970) dem onstra ted th a t the tim e for com plete d ig e s tio n

d e c re a se d for 147 hours a t 3 °C to 18 hours a t 23 °C in fingerling

so ck ey e sa lm o n . The ra te of d ig e s tio n in c re a se d w ith in c rea s in g tem ­

p era tu re from 20 °C to 23 °C in O ph iocephalus p u n c ta tu s (G erald 1973).

The p e rcen tag e of food rem aining in the stom ach of the ch an n e l c a tf ish

in c re a se d w ith d e c re a s in g w a te r tem pera tu re from 29 °C to 10 °C

(S h rab le , T iem eier, and D eyve 1969). A sadi (1967) rep o rted th a t the

s e c re tio n of d ig e s tiv e enzym es by ch an n e l c a tf ish w as low a t w a te r

tem p era tu res below 1 5 .5 ° C . Large mouth b a s s M icrop terus sa lm o ides

m ain ta in ed a t tem p era tu res of 10 , 1 5 , 2 0 , and 25 ° c req u ired 50 , 37 ,

2 4 , and 19 hours re sp e c tiv e ly fo r to ta l g a s tr ic d ig e s tio n (M olnar and

Tolg 1962).

E vidence for s im ila r tem peratu re e f fe c ts on feed in g and d i­

g e s tio n a re p re se n t in mammals (M aitrya 1961) and am ph ib ians (Teorell

1949 , Sm it 1964, B ustard 1967). K epenis and M cM anus (1974) found

th a t the tu rtle C hrysem ys p ic ta w as le th a rg ic and did no t a c c e p t food a t

15 0C . W ith an in c re a se in am bien t tem p e ra tu re , bo th food in tak e and

Page 10: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

fe c a l and u rine p roduction in c re a s e d . After fe e d in g , sp ec im en s of

P seudem ys sc rip ta in c re a se d th e ir therm al preferendum an d .b ask in g b eh av ­

io r (M oll and Legler 1971). C hem ical d ig e s tio n did no t o ccu r for two

w eek s in tu r tle s exp o sed to low tem p era tu res (Fox and M u sacch ia 1959).

M ost of th e work on p h y s io lo g ica l e ffe c ts of tem pera tu re on

sn ak es d e a ls w ith Og consum ption and h ea rt r a te . An in c re a se in body

tem p era tu re up to a ce rta in maximum is fo llow ed by an in c re a se in both

O2 consum ption and h ea rt ra te in se v e ra l s p e c ie s (Jacobsen and

W hitford 1971 , G reenw ald 1971, Buikema and Armitage 1969 , D m 'el and

Borut 1972).

- S k oczy las (1970a) found th a t d ig e s tio n in th e stom ach of the

g ra s s s n a k e , M atrix n a trix L . , ta k e s p la ce m ost rap id ly a t a tem pera­

tu re of 25 °C and may be only s lig h tly s low er a t 35 ° C . At 15 °C

d ig e s tio n in th e g ra s s snake is very slow or com plete ly in h ib ited and

n ev er o ccu rs a t 5 °C (Skoczylas 1 9 7 0 a). In the k ing s n a k e , Lam propeltis

q e tu lu s ,g a s tr ic d ig e s tio n requ ired 10 to 40 hours a t a tem peratu re of

32 ° C , about 48 hours a t 22 ° C , and d ig e s tio n did not o ccu r a t 7 °C

(Root 1961).

In c re a se d tem peratu re is known to in c re a se th re e g a s tr ic a c t iv ­

i t ie s : (1) a c tiv ity of d ig e s tiv e e n z y m e s , (2) g a s tr ic ju ic e s e c re t io n ,

and (3) g a s tr ic m otility (Smit 1967). The q u a n ti ty , pH , and p ro teo ly tic

a c tiv ity of g a s tr ic ju ic e a re a lso a ffe c te d by tem peratu re (Skoczylas

1970b).; T hus, to an optimum le v e l , in c re a s in g the tem pera tu re

Page 11: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

f a c i l i ta te s the d ig e s tiv e p ro c e s s . Above th e optimum le v e l , the tem ­

p era tu re ap p ea rs to have a re ta rd in g e ffec t in the g ra s s snake

(S koczylas 1970b) and d ig e s tio n occurred a t a s low er ra te a t 35 °C

th an a t 25 ° C .

C ow les and Bogert (1944) repo rted d ig e s tiv e fu n c tio n s in the

liz a rd Sceloporus m ag is te r may be fa c i l i ta te d by therm oregulatory

b eh av io r. E leva ted body tem p era tu res w ere recorded in a boa c o n s tr ic to r

one day a f te r feed ing by M cG innis and M oore (1969) and by K itchell

(1969). They o b serv ed th a t m ean body tem p era tu res a f te r feed ing w ere

h igher th an in unfed c o n tro ls . Regal (1966) a lso o b se rv ed th a t many

re p tile s norm ally a c tiv e a t low tem p era tu res seek h ig h er tem p era tu res

a fte r feed ing and concluded th a t the p re se n c e or a b se n c e of food in the

stom ach may be a d e term inan t in therm oregulatory b eh av io r .

A side from the o b se rv a tio n s of M cG innis and M oore (1969)

and Regal (1966), th e e ffe c t of feed ing on therm al s e le c tio n in

C o n s tric to r c o n s tr ic to r h as no t b een s tu d ie d . The p re se n t in v e s tig a ­

tio n w as undertaken to provide q u an tita tiv e da ta on fed and nonfed

s n a k e s .

Page 12: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

MATERIALS AND METHODS

C o n stric to r co n s tr ic to r from northern S in a lo a , M exico w ere

c o lle c te d in Septem ber 1973, and h e ld in 51cm x 91cm x 122cm c a g e s

u n til M ay 1 9 7 5 / The to p and one s id e of, each cag e w as co n s tru c ted of

c le a r p le x ig la s . The floor and o ther th ree s id e s w ere of p lyw ood . Each

cag e housed th re e s n a k e s . A th e rm o s ta tic a lly -c o n tro lle d e le c tr ic h e a te r

m ain ta ined tem p era tu res in th e ca g es a t 29 °C during th e lig h t period ,

and a t 26 °C during th e dark period . Light w as provided from 0600 to

2000 by overhead f lu o re sce n t l ig h ts . . W ater w as a v a ila b le ad libitum

and food w as provided a t frequen t in te rv a ls .

Therm oregulatory behav io r w as observed w hile th e snake w as

en c lo sed in a therm al g rad ien t cham ber (F ig s . 1 and 2 ). The g rad ien t

cham ber w as co n s tru c ted w ith a g a lv an ized m etal bottom and an arch ing

p lex ig las top forming the lo n g itu d in a l h a lf of a cy lin d er w ith th e d im en­

sio n s 51cm x 51cm x 198cm . The ends of th e h a lf cy lin d er w ere f itted

w ith w ooden doors through w hich the sn ak es w ere in tro d u ced to the

ch am b er. The m etal bottom w as covered by 1cm of sand and e lev a ted

above a ta b le s u rfa c e . Two 100-w att lig h t bu lbs under one end of the

cham ber provided a tem peratu re of 55 °C in th e cham ber d ire c tly above

th e b u lb s . The o p p o site end of th e cham ber w as coo led by re frig e ra te d

w ater c ircu la ted in a co iled copper tu b e . The tem peratu re on th e su rface

Page 13: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

5

©1 w ater□

a

F ig . 1. F loorp lan of g rad ien t cham ber w ith 12 u n its u sed a s p o sitio n m arkers (51cm x 51cm x 198cm ).

O 60--

3 0 -

w io

F ig . 2 . S u b stra te tem p era tu res a t e a c h p o s itio n in th e g ra d ie n t.

Page 14: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

. : ' . ' ' ' ■. of the- w a te r c o ils w as 18 °C w hile the a rea be tw een th e c o ils w as

m ain ta ined a t 20 ° C . S u b stra te tem p era tu res w ere determ ined w ith a

m ercury therm om eter in tro d u ced through sm all h o les in th e s id e s of th e

cham ber.

The s h a k e 's s tom ach tem peratu re w as determ ined by the u se o f

a b a tte ry -o p e ra te d te m p e ra tu re -se n s itiv e tra n sm itte r and an AM rad io

re c e iv e r . P u lse s from th e tra n sm itte r w ere re ce iv e d by th e rad io r e ­

c e iv e r , reco rded on a ta p e rec o rd e r , and tim ed w ith a 1 /1 0 second stop

w a tc h . The tra n sm itte r w as co n s tru c ted fo llow ing th e d e sc rip tio n of

MacKay (1970) and ca lib ra te d over a range of 20 °C to 42 °C w ith an

Ertco-76m m im m ersion m ercury therm om eter having a range of -5 °C to

105 ° C .

The tra n sm itte r w as e n c a se d in s ilic o n e and a tta c h e d to a

s trin g before feed ing to the s n a k e . The s ilic o n e e n c a se d tra n sm itte r

w as fo rc e -fe d to the nonfed a n im a l, but w as in se r te d in a fre sh ly

k ille d labo ra to ry ra t w hich w as then fed to the ex p erim en ta l snake to

o b ta in tem p era tu res a fte r e a tin g . The tra n sm itte r w as re tr ie v e d a t th e

co n c lu s io n of each experim ent by w ithd raw al of the a tta c h e d s tr in g .

Stom ach tem pera tu re m easurem ents w ere reco rd ed a t 3 - hour

in te rv a ls beg inn ing a t the tim e the tra n sm itte r w as in tro d u ced in to th e

sn a k e . Fed (tran sm itte r in a ra t c a rc a s s ) and nonfed con tro l tem pera­

tu re m easu rem en ts w ere in it ia te d a t 1 2 00 , 1700, and 2200 , and

co n tin u ed over a 9 6 -h o u r p e rio d . A l e a s t sq u ares p e rio d ic re g re ss io n

Page 15: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

lin e w as draw n through the d a ta p o in ts . A nalysis of v a rian ce u s in g an

f t e s t w ith a random ized b lock d es ig n w as u se d to determ ine le v e ls of- . x .

s ig n if ic a n c e .

Page 16: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

RESULTS

Stom ach tem p era tu res s e le c te d over a 96-hour t e s t period are

p re sen ted in F igures 3 through 20. Therm al s e le c tio n in C o n s tric to r

c o n s tr ic to r ap p ears to be of a cy c lic n a tu re w ith c y c le s approxim ating

24 h o u rs „ The h ig h e s t tem p era tu res w ere s e le c te d in th e .la te ev en in g ;

b e tw een a few hours prior to s u n se t and approx im ate ly m id n ig h t. The

lo w e s t tem p era tu res w ere se le c te d be tw een the ea rly m orning hours and

n o o n . R ecordings of stom ach p o s itio n and tem peratu re re la te the tem ­

perature f lu c tu a tio n s to the behav io r of the sn ak es who a c tiv e ly change

th e ir p o s itio n in th e therm al g rad ien t (F ig s . 3 -2 0 , p ag es 1 1 -1 9 ).

The f ir s t m ajor a c tiv ity period ap peared a fte r a low tem peratu re

w as rea ch e d in th e e a rly to la te morning w hen the sn ak e moved tow ard

th e warm end of th e g ra d ien t. The seco n d m ajor a c tiv ity period u su a lly

occurred betw een m idnight and dawn when th e snak e moved tow ard the

co o ler end of th e g rad ie n t. T hese o b se rv a tio n s ag ree w ith M cG innis and

M bore (1969), who found the a c tiv ity of C o n stric to r c o n s tr ic to r during

a 24-hour period to be bim odal w ith m ajor m ovem ents occu rring about

noon and about m idnigh t.i '

M ean s e le c te d stom ach tem p era tu res are p re sen ted in Table 1.

F eeding re su lte d in a s ig n if ic a n t in c re a se in m ean s e le c te d tem p era tu re ,

bu t did not a lte r th e b a s ic d iurnal cy c le observed in nonfed s n a k e s . Fed

8

Page 17: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

9

Table 1. M ean S e lec ted Stom ach T em peratures ° C .

F eed ing Snake 1200Time of F eed ing

1700 2200

#1 2 9 .1 2 9 .2 . 3 1 .1

Fed ■ . #2. 2 6 .8 3 1 .3 3 0 .7

#3 3 1 .7 2 9 .9 3 0 .8

. #1 2 8 .2 2 5 .7 - 2 7 .0

Nonfed #2 2 5 .5 2 6 .1 2 5 .6

#3 2 7 .4 2 7 .2 2 8 .5

Table 2. V oluntary M aximum T em peratures ° P .

Feeding Snake 120.0Time of F eed ing

1700 2200

#1 3 6 .6 3 6 .9 3 6 .1

Fed . #2 3 5 .0 3 6 .4 3 5 .2

#3 3 6 .2 3 6 .0 3 5 .1

#1 3 5 .2 3 6 .1 3 2 .0

NOnfed #2 3 0 .0 3 3 .8 3 3 .8

' #3 3 3 .9 3 1 .2 3 1 .2

Page 18: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

■ ::: . ■ : 10

sn ak es b eh av lo ra lly m ain ta ined more e le v a te d te m p e ra tu re s . D iffe rences

in m ean tem perature s e le c t io n due to th e time of feed in g w ere not s ig n if ­

ic a n t . . ;

Voluntary therm al maximum a s defined by C ow les an d Bogert

(1944) w as h igher in fed sn a k e s than in nonfed an im als (Table 2). The

h ig h e s t s tom ach tem perature recorded w as 3 6 .9 °G in a fed sn a k e . The

mean vo lun tary therm al maximum for the th ree fed s n a k e s w as 3 6 .5 ° C ,

and for the th ree nonfed sn ak es w as 3 4 .6 °C . Brattstrom (1965) reported

a vo lun tary therm al maximum of 34 ° C Z and M cG inn is and M oore (1969)

recorded a maximum of 3 8 .5 0C for a spec im en of C o n s tr ic to r c o n s t r ic to r .

The la t te r au thors observed the 3 8 .5 °C read ing only o n c e , and i t w as

th e ir only record ing above 30 ° C .

Voluntary minimum tem pera tu res of 2 1 .3 °C and 2 0 .1 °C w ere

recorded for a fed and a nonfed sn a k e , r e s p e c t iv e ly . There were no

s ig n if ican t d iffe ren ces be tw een fed and nonfed sn ak es in th is r e s p e c t .

Page 19: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

Figures 3> 5 , 7 , 9 , 11, 13, 15, 17 , 19: (Selected Stomach Temperatures) .

.F igures 4 , 6 , 8 , 10 , 12 , 14 , 16 , 18 , 20: (Selected Stomach P o s i t io n s ) ,.

© — Observed, pos ition points for fed snake

O bserved pos ition poin ts for nonfed ^ *"* snake.i- • '

. (See F i g . 1 and Fig. 2 for su b s t ra te tempera tures a t e ach p o s i t io n . )

L east sq u a res periodic re g re s s io n line for fed snake

L east squares period ic re g re s s io n line for nonfed snake

M ean tem pera tu res for fed snake

M ean tem pera tu res for nonfed snake

^/ / — Dark period (2000 - 0600)

O bserved da ta points for fed snake

O - O bserved da ta poin ts for nonfed . snake

Page 20: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

11

36+

-2 9 .1-2 8 .2

• s

CZD,2 4 --

22 - -

Hours after FeedingFig . 3. S e lec ted s tom ach tem pera tu res for boa #1 fed a t 1200.

12- -

# i

S 8 -

# # #

E 6 - -

o o 00

O O< O

Hours after Feeding

Fig. 4. S e lec ted stom ach pos it io n s for boa #1 fed a t 1200.

Page 21: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

1 2

3 64

34--

32--

5 3 0 - ,

E 28--

26--V -

/*C/D

24--

22 -

u L

Hours after FeedingFig. 5. S e lec ted stom ach tem pera tu res for boa #2 fed a t 1200.

1 2 -

10- - #

i »# i

Hours a fter FeedingFig. 6. Se lec ted s tomach pos i t ions for boa #2 fed a t 1200.

Page 22: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

13

36-

A

•31.7

E 2 8 -f •27.4

2 6 -

C/D24--

2 2 -

Hours after FeedingF ig . 7. S e lec ted stom ach tem pera tu res for boa #3 fed a t 1200.

12-

6 6g in-- s • • • « # • O• e OO o

#

OO

o 4_.

o o o'<> o o

Hours after Feeding

Fig. 8. Se lec ted stomach pos i t ions for boa #3 fed a t 1200.

Page 23: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

14

36 + oo

34--

32--

B 30i-•29.2

to 26- i '----- 25.7/ “

uD2 4 -

22-

Hours after FeedingFig . 9. S e lec ted s tom ach tem pera tu res for boa #1 fed a t 1700

oo10-

• • •• o

##

O DO o o or o o go o

o o o

Hours a f te r FeedingFig. 10. S e lec ted s tomach pos i t ions for boa #1 fed a t 1700.

Page 24: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

15

36--

o #

-3U

E 28--

26.1

CO

22- -

Hours after Feeding Fig . 11. S e lec ted s tom ach tem pera tu res for boa #2 fed a t 1700.

8-

0 o§ 6~ *

©oo o

Hours af te r FeedingFig. 12. Se lec ted s tomach pos i t ions for boa #2 fed a t 1700.

Page 25: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

364-

34--

•31.2

— y------------\ --------- 07— v — /---- 27.2— r 10fD1■M

26--

co2 4 --

22 -

Hours after FeedingFig. 13. S e lec ted s tom ach tem pera tu res for boa #3 fed a t 1700.

12- -

4->

§-o 10--,.##fe, "5 8 - - « > 0•6 ° fOo 6”+-»(0

» • •«• #

C O O

• #

:o■Mto

£ O O

Hours after FeedingFig. 14. Se lec ted s tomach pos i t ions for boa #3 fed a t 1700.

Page 26: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

36+

# #

32--

CO 062 4 --

2 2 -

ezzHours after Feeding

Fig. 15. S e lec ted stom ach tem pera tu res for boa #1 fed a t 2200.

1 2 -

o<#e

• • s> 0

o oE 6 - -

oo06 0

Hours after Feeding

Fig. 16. S e lec ted stom ach po s it io n s for boa #1 fed a t 2200.

Page 27: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

18

30.730--

t - i

--A - "t 25.6CO

2 2 -

N , , W , , W , , W" F70 6 12 18 24 30 36 42 48 54 60 66 72 78 84 90 96

Hours a fter Feeding Figure 17. S e lec ted stom ach tem pera tu res for boa #2 fed a t 2200.

1 2 - • •cQ)

&

10 - - • I • •• ee

• ex:oro§+j

• IWO§a

Hours after FeedingFig. 18. Se lec ted stomach p os i t ions for boa #2 fed at 2200.

Page 28: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

19

32--•30.8

— 28.5E 2 8 --

2 6 --

co 24--

22- -

m , ,n72 78 84 90 960 6 12 18 24 30 36 42 48 54 60 66 : :

Hours a fter FeedingFig. 19. S e lec ted stom ach tem pera tu res for boa #3 fed a t 2200.

12- -

I mi£Cn o c 8 - -

o • • • # • • O# ## *

# #

o o o0fD1■M o o o03

4-1o§03

£

Hours a f te r Feed ingFig. 20. Se lec ted s tomach pos i t ions for boa #3 a t 2200.

Page 29: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

DISCUSSION AND CONCLUSIONS

The in c re a se in both maximum volun tary and mean preferred

tem perature in fed sn ak es w as c o n s is te n t w ith Regal.'s (1966) o b se rv a ­

tio n th a t C o n s tr ic to r co n s tr ic to r w as more therm ophilic a f te r feeding

than befo re . This therm ophilia is s im ila r to th a t found in tu r t le s

(Gatten 1974, Moll and Legle r 1971). H ow ever, r e s u l t s from Caiman

c rocod ilus in d ica te th a t preferred tem pera tu res are in d ep en d en t of time

s in ce l a s t feed ing (Diefenbach 1975). The h igher s tom ach tem pera tu res

s e le c te d compared w ith Sm it 's (1967) finding th a t d ig e s t iv e p ro c e s s e s

are more e ff ic ien t a t a tem perature s lig h tly h igher th an the normal m ean

body tem perature in f i s h . H ydrochloric a c id co n cen tra tio n in the g a s t r ic

ju ice and p ro teo ly tic a c t iv i ty in the s tom ach proceed f a s te r a t e le v a te d

body tem pera tu res (D iefenbach 1975, S koczy las 1 9 7 0 a , and Asadi 1967).

E levated stom ach tem pera tu res cou ld be a c t in g to in c re a se

blood flow to the g a s tr ic g lands d e liver ing more oxygen and nu trien ts to

c e l l s producing the g a s tr ic se c re t io n s a s in frogs (Smit 1964). Lucey

(1974) found th a t heating of the sk in or c a rap ace of the tu r t le s P .

f loridana and C . se rpen tina w as fo llow ed by an in c re a se in blood flow

to the h ea ted a r e a . An in c re a se in blood flow to the s tom ach brought

abou t an in c re a se in a c id sec re t io n in d o g s '(P e te r e t a l . 1962). An

in terrup tion of the blood supply to the stom ach of the in ta c t dog rap id ly

‘ ■ ' . ■ ■ ■ ■ 2 0 ■ v : : v -

Page 30: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

in d u ces a c e s s a t io n of sec re t io n (Coy e t ' a l . 1951). The in c re a se d

tem pera tu res would a lso exert a d irec t e ffec t a c c e le ra t in g the m etabo­

lism of the g a s tr ic g land c e l l s and thereby in c re a s in g th e ir production

of a c id and p ro teo ly tic enzym es (Teorell 1949). In c re a s e s in s tom ach

m otility due to in c r e a s e s in tem perature would a lso a c c e le r a te g a s tr ic

s e c re t io n s (Smit 1967). At low am bient te m p e ra tu re s , s tom ach HC1

se c re t io n and p ro teo ly tic a c t iv i ty are a t a minimum or to ta l ly la c k in g ,

thus inh ib iting g a s tr ic d ig es t io n (Skoczylas 1970a, A sadi 1967).

Optimum d ig e s t iv e a c t iv i ty in C o n s tr ic to r co n s tr ic to r ap p eared not to

follow p a s s iv e ly , but com pelled the anim al to seek h igher am bient

tem pera tu res to f a c i l i ta te d ig e s t io n .

The mean preferred body tem perature s e le c te d by C onstric to r

c o n s tr ic to r in th is s tudy of 2.6.8 °C can be compared to 30 °C reported

by Brattstrom (1965). M cG innis and Moore (1969) show ed th a t juven ile ,

boa c o n s tr ic to rs had mean preferred body tem pera tu res of around 30 °C

in outdoor e n c lo su re s in San B ia s , M ex ico , The mean preferred tem ­

pera tu res of th ree a d u lts in a phototherm al cham ber w ere 2 2 .6 , 23 .1 ,

and 2 3 .7 . The th re e -d a y mean body tem perature of a f re e - l iv in g adu lt

boa m easured by rad io -te lem etry w a s 2 6 .4 . Carpet py thons M orelia

sp i lo te s have a preferred body tem perature of 30 °C in th e f ie ld and

in confinem ent (Webb and H eatw ole 1971). Cogger and Holm es (1960)

reported a preferred body tem perature for th is s p e c ie s of 34 ° C . S pec i­

mens of Boa co n s tr ic to r found in the f ie ld in Sonora, M exico had

Page 31: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

2 2

body tem pera tu res ranging from 2 5 .4 °C to 3 0 .4 0 C (Myres and Ells

1968).

, Although a con tinuous hea t source w as a v a i la b le , the an im als

vo lun ta ri ly ch o se to vary th e ir tem perature ra ther th an rem ain a t a

c o n s ta n t e le v a te d body tem pera tu re . This supports Regal.'s (1966)

finding th a t the l iza rd Klauberina r iv e rs ia n a may prefer ra ther than

simply to le ra te low environm ental te m p e ra tu re s . Regal (1966) reported

a minimum body tem perature for the boa co n s tr ic to r of 31 °C a f te r

feeding . My re s u l t s in d ica te a much low er minimum tem pera tu re a f te r

f e e d in g , often below 26 ° C .

The d iurnal tem perature s e le c t io n by C o n s tr ic to r co n s tr ic to r i s

b e s t co rre la ted with ligh t and dark p e r io d s , ra ther th an time of fe e d in g .

The h ig h es t tem peratures s e le c te d and the in c re a se in th e s n a k e 's

a c t iv i ty during the la te evening and n igh t support the b e l ie f th a t

C o n s tr ic to r co n s tr ic to r is primarily a noc tu rna l s p e c ie s (McGinnis and

Moore 1969).

Page 32: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

REFERENCES

A s a d i , S. 1967. Effect of tem perature on the d ig e s t iv e enzym es of channe l c a t f ish Ic ta lu ru s punc ta tu s (R afinesque) . M .S . t h e s i s , K ansas S tate U n i v . , M an h a ttan , K a n s a s .

Brattstrom , B. H . 1965. Body tem pera ture of r e p t i l e s . Am. M idi.N a t . 73: 3 76 -422 .

Brett, J . R. and H ig g s , D . A. 1970. Effects of tem pera ture on the ra te of g a s t r ic d ig es t io n in f ingerling sockeye sa lm on , O nchorhynchus n e rk a . I . F ish R es . Bd. C an . 27 : 1767-1779 .

Buikema, A. L. and Armitage, K. B. 1969. The e ffec t of tem perature on the m etabolism of the pra irie ringneck s n a k e , D iadophis punc ta tu s a ry n i . H erpeto log ica 25 : 194 -206 .

B ustard , H. R. 1967. Activity cyc le and therm oregulation in the A ustra lian gecko Gehyra v a r i e g a ta . Copeia 4: 753-758 .

C ogger, H. G . and H o lm es , A. 1960 . Thermoregulatory behavior in a specim en of M orelia sp i lu te s v a r ieg a ta Gray (Serpentes:

• Boidae). P roc . Linn Soc . New So. W a les 8 5 : 3 2 8 -3 3 3 .

C o w le s , R. B. and BOgert, C . M. 1944. A prelim inary study of the therm al requirem ents of d e s e r t r e p t i l e s . Bull. Amer. Mu s . N a t . H is t . 83: 267-296..

C oy , F . E . ; B u jandas , F. J . ; D eG raffen re id , T .P . ; and Rehm, W . S.1951. Effect of in terrupting blood flow on g a s t r ic po ten tia l and HC1 s e c re t io n . G as troen tero logy 17 : 2 6 0 -2 6 8 .

D ie fen b ach , C . O . 1975. Thermal p re fe ren ces and therm oregula tion• in Caiman c ro c o d i lu s . C opeia 2 ; 102-123 .

D m 'e l, R. and Borut, A. 1972. Thermal b ehav io r , h e a t ex ch an g e , and m etabolism in the d e se r t snake S palerosoph is c l i f fo rd i i . P h y s io l . Zool. 4 5 : 7 8 -9 4 .

Fox, U . M . and M u sa c c h ia , X. J. 1959.. N otes on th e pH of the d ig e s t iv e t ra c t of Chrysem ys p i c t a . Copeia 4: 3 3 7 -3 3 9 .

23

Page 33: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

24

G a t te n , R„ E. 1974. Effect of nu tri t iona l s ta tu s on the preferred body tem perature of th e tu r t le s Pseudem ys sc r ip ta and Terrapene o m a t a . C opeia 4: 619-648 . .

G era ld , V. M . 1973. Rate of d ig es t io n in O phiocephalus punc ta tus Bloch. Comp. Biochem. P h y s io l . 4 6 : 1 9 5 -2 0 5 .

G reenw ald , O. E. 1971. The e ffec t of body tem perature on oxygen consum ption and heart ra te in the Sonoran gopher sn a k e , P ituophis ca ten ife r aff in is H a llo w ell . C opeia 1: 9 8 -1 0 6 .

Ja c o b se n , E. R. and W h itfo rd , W . G . 1971. P h y s io lo g ica l r e sp o n se s to tem perature in the p a tc h -n o se d s n a k e , Salvadora h exale p i s . H erpeto log ica 27: 28 9 -2 9 5 .

K e p e n is , V. and M cM anus , J. 1974. B ioenergetics of young pa in ted tu r t le s Chrysem ys p i c t a . Comp. Biochem. P h y s io l . 4 0 : 3 0 9 -3 1 7 .

K itche ll , J. F . 1969. Thermophilic and therm ophobic r e sp o n se of sn ak es in a therm al g ra d ie n t . C opeia 1: 1 8 9 -1 9 1 .

L u c ey , E„ C . 1974. H eart ra te and p h y s io lo g ica l therm oregulation ina bask ing tu r t le , Pseudem ys Scripta E le g a n s . Comp. Biochem. P h y s io l . 48: 4 7 1 -4 8 2 .

MacKay, R. S . 1970. B io -M edica l T e lem etry . Second ed i t io n . John W iley and S ons, New York, N . Y.

M a it ry a , B. B. 1961. Effect of tem pera tu re on g a s t r ic s e c re t io n .J. S c i . In d u s tr . R es. 2 0 : 3 5 5 -3 5 6 .

M cG in n is , S. M. and M oore, R. G . 1969. Thermoregulation in the Boa C o n s tr ic to r , Boa c o n s t r ic to r . H erpe te log ica 25 : 3 8 -4 5 .

M oll, E. O . and L eg le r, J . M. 1971. The life h is to ry of a neo trop ica l s l id e r t u r t l e , Pseudem ys sc r ip ta in P an am a. Bull. L. A. Co.. M u s . N a t . H is t . Sc i. 11 : 1 -1 0 2 .

M olnar, G . and Tolg, I . 1962. Relation be tw een w a te r tem perature and g a s tr ic d ig e s t io n of large mouth b a s s M icrupterus sa lm oides L a c e p e d e . T. F ish . R es . Bd.' C a n . 1 9 : 10 0 5 -1 0 1 2 .

My r e s , B. C . and E l ls , M. M. 1968. Thermal agg reg a tio n in Boa c o n s t r ic to r . H erpeto log ica 24 : 6 1 -6 8 .

Page 34: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature

25

P e te r , E. T . ; N ico lo ff , D. M . ; S o s in , H . ; W alder , A. I . ; andW a g e n s te e n , O. H. 1962. R e la tionsh ips be tw een g a s tr ic blood flow and s e c re t io n . Fed . Proc . 2 1 : 264 .

Regal, P . J . 1966. Thermophilic r e sp o n s e s following feeding in ce r ta in r e p t i l e s . Copeia 3: 588 -5 9 0 .

Root, H . D. 1961. G a s tr ic d ig es t io n w ith hypothermia: O b serva tions and a p p l ic a t io n s . T h e s is , U n ivers ity of M inneso ta .

Shrable , J. B.; T iem eier, O. W . ; and Deyve , C . W . 1969. Effects of tem perature on rate of d ig e s t io n by channel c a t f i s h . Progr. F ish . C u lt . 31: 131 -138 .

S k o c z y la s , R. 1 9 7 0 a . In fluence of tem perature on g a s t r ic d ig es t io nin the g ra s s s n a k e , Natrix na tr ix L . Comp. Biochem. P h y s io l . 33: 7 93 -804 .

________. 1970b. Salivary and g a s tr ic ju ice sec re t io n in the g ra sss n a k e , N atrix natr ix L. Comp. Biochem. P h y s io l . 35 : 885-903 .

Sm it, H. 1964. The regu la tion of p ep s in sec re t io n in the ed ib le frog Rana e sc u le n ta L. Comp. Biochem. P h y s io l . 13 : 129-141 .

________. 1967. In fluence of tem perature on the ra te of g a s tr ic ju icese c re t io n in the brown bullhead Ic ta lu rus n e g u lo s u s . Com p. Biochem. P h y s io l . 21 : 125-132 .

T eore ll , T. 1949. On the in fluence of tem perature on the HC1 form a­tion and the oxygen consum ption of the s to m a c h . Experientia 5: 40 9 -4 1 0 .

W ebb, G. and H e a tw o le , H . 1971. P a tte rns of hea t d is tr ib u tio n w ith in the bod ies of some A ustralian p y th o n s . C opeia 2: 209-220 .

Page 35: EFFECTS OF FEEDING ON THERMAL SELECTION …arizona.openrepository.com/arizona/bitstream/10150/347972/1/AZU_TD...EFFECTS OF FEEDING ON THERMAL SELECTION BY CONSTRICTOR ... perature