First published online June 15, 2006
Journal of Experimental Biology 209, 2495-2508 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02294
Metabolic organization of freshwater, euryhaline, and marine elasmobranchs: implications for the evolution of energy metabolism in sharks and rays
B. Speers-Roesch1,
Y. K. Ip2 and
J. S. Ballantyne1,*
1 Department of Integrative Biology, University of Guelph, Guelph, Ontario,
NIG 2W1, Canada
2 Department of Biological Science, National University of Singapore, Kent
Ridge, Singapore 117543, Republic of Singapore

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Fig. 1. Carnitine palmitoyl transferase (CPT) activities (mean ± s.e.m.) in
liver and extrahepatic tissues of freshwater, euryhaline and marine
elasmobranchs with varying levels of urea: Potamotrygon motoro
(freshwater) (no urea), Himantura signifer in freshwater (low urea),
H. signifer acclimated to half-strength seawater ( Seawater)
(increased urea), Taeniura lymma (marine) (high urea), and
Chiloscyllium punctatum (marine) (high urea). See text or
Fig. 4 for specific urea
levels. Sample sizes and the results of statistical comparisons between
tissues and species are given in Tables
1 and
2. The salinity (and thus urea
level) for each species or acclimation group is indicated by the shading of
the bars. Enzyme measurements were made at 25°C. ND, not detectable.
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Fig. 2. 3-Hydroxyacyl CoA dehydrogenase (HOAD) activities (mean ± s.e.m.) in
liver and extrahepatic tissues of freshwater, euryhaline and marine
elasmobranchs with varying levels of urea: Potamotrygon motoro
(freshwater) (no urea), Himantura signifer in freshwater (low urea),
H. signifer acclimated to half-strength seawater ( Seawater)
(increased urea), Taeniura lymma (marine) (high urea) and
Chiloscyllium punctatum (marine) (high urea). See text or
Fig. 4 for specific urea
levels. Sample sizes and the results of statistical comparisons between
tissues and species are given in Tables
1 and
2. The salinity (and thus urea
level) for each species or acclimation group is indicated by the shading of
the bars. Enzyme measurements were made at 25°C. ND, not detectable.
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Fig. 3. D-ß-hydroxybutyrate dehydrogenase (D-ß-HBDH)
activities (mean ± s.e.m.) in liver and extrahepatic tissues of
freshwater, euryhaline and marine elasmobranchs with varying levels of urea:
Potamotrygon motoro (freshwater) (no urea), Himantura
signifer in freshwater (low urea), H. signifer acclimated to
half-strength seawater ( Seawater) (increased urea), Taeniura
lymma (marine) (high urea) and Chiloscyllium punctatum (marine)
(high urea). See text or Fig. 4
for specific urea levels. Sample sizes and the results of statistical
comparisons between tissues and species are given in Tables
1 and
2. The salinity (and thus urea
level) for each species or acclimation group is indicated by the shading of
the bars. Enzyme measurements were made at 25°C. ND, not detectable.
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Fig. 4. Relationship between white muscle urea concentration and liver glutamate
dehydrogenase (GDH) activity in elasmobranchs. Values are mean ± s.e.m.
See text for full names of species. Error bars for GDH in Potamotrygon
motoro and P. magdalenae are asymmetrical to avoid overlapping
bars. 1GDH values are from the present study (see
Table 1 for sample sizes) and
urea values are from the same animals (N=58, taken from
various studies) (Treberg et al.,
2006 ). 2GDH value
(Singer and Ballantyne, 1989 );
urea value was assumed to be the same as in P. motoro
(Treberg et al., 2006 ).
3GDH value (Moon and Mommsen,
1987 ); urea value (Forster and
Goldstein, 1976 ). 4GDH value
(Battersby et al., 1996 ); urea
value (Treberg and Driedzic,
2002 ). GDH activities for spiny dogfish and little skate were
adjusted to 25°C using Q10=2. The regression is significant
(r=0.93, P<0.001, linear regression ANOVA).
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© The Company of Biologists Ltd 2006