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First published online April 18, 2006
Journal of Experimental Biology 209, 1651-1661 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02195
Metabolic scaling associated with unusual size changes during larval development of the frog, Pseudis paradoxus
Departamento de Fisiologia, Instituto de Biociências, Universidade de São Paulo, Rua do Matão, Travessa 14, no 321, Cidade Universitária, CEP 05508-900, São Paulo, SP, Brazil
* Author for correspondence (e-mail: scrsouza{at}ib.usp.br)
Accepted 2 March 2006
The early larvae of P. paradoxus grow large but metamorphose into
relatively small frogs, the diminished post-metamorphic growth producing a
marked contrast between maximum larval size and adult. Thus, O2
uptake does not appear to limit the energy expenditure on growth processes,
and unlike in other anuran larvae, may not be a surface area-related function
in P. paradoxus larvae. The resting rates of metabolism
(
O2) and
partitioning between aquatic
(
wO2) and aerial
O2 uptake
(
aO2) were
measured on tadpoles and froglets by closed system respirometry, using water
of PO2 ranging from 145 to 40 mmHg. Correlative
changes in body glycogen and lactate were examined by standard enzyme assays.
Scaling patterns in the growth and degrowth stages were analysed on
whole-body, log-transformed data using linear regressions. In normoxia,
O2 was
2.12.5 µmol g1 h1 in the early
larvae, increasing more than twofold on forelimb emergence and decreasing
sharply in the froglets;
O2 varies in
strict proportion to body mass (Mb), both in the growth
(b=1.02) and degrowth (b=0.97) phases, according to the
equation
O2=aMbb,
where b is the scaling coefficient.
wO2 constitutes
>90% of total uptake in the growth stages, increasing with b=1.02
while
aO2
increases with b=1.13; during degrowth there is a change in the
pattern related to intensification of metamorphosis. Hypoxic water did not
affect
O2;
however, in all larval stages
wO2 and
aO2 changed with
a decrease in PO2. At 60 mmHg, rates are more
severely affected in the largest tadpoles, causing the b values for
wO2 and
aO2 to change to
0.11 and 1.44, respectively, in the growth phase. Glycogen and lactate levels
increase out of proportion with body mass increase (b=2.05 and 1.47,
respectively) in the growth stages, and increase anaerobic capacity in late
metamorphosis. In hypoxic water, glycogen levels decrease in the growth stages
and the largest tadpoles accumulate surplus lactate, possibly related to
surfacing activity. Our results may reveal the consequences of size on energy
demand at the tissue level in P. paradoxus larvae, indicating that
air breathing must subsidise energy expenditure during larval development.
Key words: metabolic rate, oxygen uptake, metamorphosis, scaling, hypoxia, glycogen, lactate, Amphibia, anuran, Pseudis paradoxus