First published online June 15, 2006
Journal of Experimental Biology 209, 2554-2566 (2006)
Published by The Company of Biologists 2006
doi: 10.1242/jeb.02259
Following the heart: temperature and salinity effects on heart rate in native and invasive species of blue mussels (genus Mytilus)
Caren E. Braby* and
George N. Somero
Hopkins Marine Station, Department of Biological Sciences, Stanford
University, Oceanview Boulevard, Pacific Grove, CA 93950, USA

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Fig. 1. Heart rate response to temperature and methods for impedance pneumography.
Heart activity patterns from a single individual over a range of temperatures
from 14.531°C are shown. Each heart rate trace spans 3 min. Inset
shows the schematic of the heart rate monitoring apparatus, and the placement
of electrodes. Mussel drawing adapted from
(Brusca and Brusca, 1990 ).
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Fig. 2. Environmental exposure and heart rate response to stress in M.
trossulus. (A) Heat stress; as temperature increases (broken line;
beginning at 01:00 h, experimental time), heart rate increases until the
animal reaches a temperature at which heart rate falls rapidly
(Hcrit). With continued heat stress, heart rate remains
depressed, but as soon as temperature drops, heart rate returns to a normal
level. (B) Cold stress; as temperature decreases from acclimation level, heart
rate decreases to zero. (C) Low salinity stress; as salinity decreases from
acclimation level, heart rate decreases very gradually until the animal
reaches a salinity at which there is a rapid fall in heart rate
(Scrit). Heart rate remains depressed during continued low
salinity treatment but recovers immediately when salinity begins to
increase.
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Fig. 3. Heart rate response to temperature and salinity in differently acclimated
mussels. Species are referred to by the first letter of the species' name:
M. trossulus (t), M. galloprovincialis (g), M.
edulis (e), and M. trossulus/M. galloprovincialis
hybrids (h). Symbols connected by solid lines are 14°C-acclimated; symbols
connected by dotted lines are 21°C-acclimated. Values are means ±
s.e.m. For N and P values, see
Table 2. (A) Resting heart rate
measured at the acclimation temperatures (14°C or 21°C). (B) Response
to heat stress (Hcrit); all species show an increase in
Hcrit with increased acclimation temperature. While there
are some differences in Hcrit among salinity treatments,
salinity acclimation does not contribute statistically to the observed
pattern. (C) Response to cold stress (lowest heart rate at 0°C); at
comparable acclimations, M. trossulus has an equal or higher heart
rate than M. galloprovincialis. Note that to make overlapping symbols
visible, we offset the g-14°C data by +0.2°C along the
x-axis. (D) Response to low salinity stress
(Scrit); there is a clear increase in
Scrit with increasing salinity acclimation in all
species.
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Fig. 4. Temperature compensation of resting heart rate (RHR; beats
min1). t, M. trossulus; g, M.
galloprovincialis; e, M. edulis. Symbols connected by solid
lines are observed RHR values averaged across salinity treatments. Symbols
connected by broken lines are observed RHRs at 14°C and expected RHRs at
21°C, using the species-specific Q10 estimate and assuming no
physiological compensation. If there was complete temperature compensation,
there would be no difference between the observed values for 14°C- and
21°C-acclimated animals (symbols connected by a horizontal line; not
shown). However, the observed RHR at 21°C is lower than expected for all
species, indicating partial compensation with acclimation temperature.
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Fig. 5. Mortality in temperature and salinity acclimation treatment groups. There
was nearly equal mortality of M. edulis (e) and M. trossulus
(t) in 2003, regardless of acclimation treatment, and both species had greater
mortality at the warmer acclimation temperature. In the following year, M.
trossulus had equivalent mortality at the 14°C acclimation and
decreased mortality under 7°C acclimation, reinforcing the positive
relationship between acclimation temperature and mortality. Only three M.
galloprovincialis (g) individuals died in both acclimation years (out of
348 total), and all three were from the 21°C acclimation in 2003 (M.
galloprovincialis 2004 is not included in the graph because there was no
observed mortality).
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© The Company of Biologists Ltd 2006