J. therm. Biol. Vol. 16, No. 6, pp. 33%343, 1991 0306-4565/91 $3.1)0+ 0.00
Printed in Great Britain. All rights reserved Copyright © 1991 Pergamon Press plc
THERMAL RESPONSES OF THE FRESH WATER TURTLE
MAUREMYS CASPICA TO STEP-FUNCTION CHANGES IN
THE AMBIENT TEMPERATURE
T. PAGES,J. F. FUSTERand L. PALACIOS
Unidad de Fisiologia, Departamento de Bioquimica y Fisiologia, Universidad de Barcelona,
Avenida Diagonal, 645 E-08071 Barcelona, Spain
(Received 18 August 1990; accepted in revisedform 3 August 1991)
Abstract--1. The turtle Mauremys caspica cools significantly faster than it heats in air. The heating/cooling
ratio is 0.49.
2. The variation of body temperature in relation to time-course in response to a step-function change
of environmental temperature, fitted to a second-order system improves that of a first-order system.
3. The gradient between ambient temperature (Ta) and equilibrium body temperature (Tb) increases
significantly and progressively when ambient temperature rises over 25°C.
4. At 40°C thermoregnlatory hyperventilation was detected, implying an increase in air convection
requirement (ventilation relative to 02 consumption, l;'E/l?o2).
Key Word Index: Turtles; heating and cooling rates; thermoregulation; ventilation; Mauremys caspica;
reptile
INTRODUCTION
Turtles are ectotherms and as such rely primarily on
heat from the environment to achieve and maintain
body temperature within the intervals necessary for
their daily activities. The primary means of thermo-
regulation in turtles, as in all ectotherms, is be-
havioural, and they simply avoid extremely hot
environments. When denied behavioural means, or
when behavioural means are not sufficient to combat
heat stress in the environment, some turtles can use
physiological mechanisms of control to keep body
temperatures (T b) below lethal limits for short
periods of time.
Turtles raise their Tb through the absorption of
solar radiation by basking in water or air (Boyer,
1965; Spotila et al., 1984). Basking is a complex
behavioural pattern affected by nutritional status,
acclimation, season and sex of the individual
(Hammond et al., 1988). When ambient temperature
(Ta) is below the preferred Tb the turtles can seek a
step change in Ta by means of basking behaviour or
water immersion.
Some studies carried out in turtles on heat transfer
relationships show different strategies for thermal
control, such as behavioural mechanisms (Sturbaum
and Riedesel, 1974; Meek, 1984; Jading et al.,
1984, 1989) and physiological thermoregulatory
systems (Weathers, 1970; Weathers and White,
1971; Cloudsley-Thompson, 1974) that differ in their
thermal dynamics from a model in which heat trans-
fer is based on thermal conductivity. Sturbaum (1982)
has reviewed the main thermoregulatory responses in
turtles, with a description of the different physiologi-
cal mechanisms used to avoid critical temperatures
and maintain body temperature within their thermo-
preferendum. This author suggested a role for respir-
atory responses to hyperthermia. Bartholomew
(1982) and Turner (1982) agree with this approach
and also suggest a possible control of body tempera-
ture by means of vasomotor responses with adjust-
ments of blood flow within the body.
Most of the studies on thermoregulatory responses
in turtles have been carried out on North American
species. It would also be interesting to analyse ther-
mal responses in a turtle from the Mediterranean
area, Mauremys caspica. Few studies have been car-
ried out on this species, and in particular, studies on
physiological thermoregulatory mechanisms in this
species have not been reported. In this study, we
determined the rate of change of body temperature
following exposure of animals to step changes of
ambient temperature. The step changes simulate ther-
mal alterations which the animal experiences under
natural conditions in basking. Analysis of the dy-
namic characteristics of these processes may allow us
to predict other patterns of thermal variation and
indicates that a ventilatory response to hyperthermia
has a regulatory role.
MATERIALS AND METHODS
Twenty-one fresh water turtles, M. caspica, with
body mass between 75 and 600 g, were used. The
animals were caught in Laguna de Adra (Almerla,
southeastern Spain).
Before experimental studies began, the animals
were maintained for 2 weeks under laboratory con-
ditions at 20-25°C, without food but with water ad
libitum. Body mass was determined before acclim-
ation. Experimental ambient temperatures were
achieved by means of two thermostatic chambers.
During the heating phase, the chamber was con-
trolled at 30 _+ I°C to represent a physiological tem-
perature and until 47 ± 2°C for extreme temperature
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