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rabbit (or human ventricle), and this drives rapid Ca 2 þ extrusion via Na þ /Ca 2 þ
exchange at a time when [Ca 2 þ ] i is very high ( Fig. 6 B). In the rabbit ventricle, the
longer action potential plateau keeps Na þ /Ca 2 þ exchange in check, delaying
extrusion until a later time where [Ca 2 þ ] i
is lower. Another implication of
Fig. 5 B is that there is net Ca 2 þ e
Z
ux during the contraction in rat (vs. net influx
in rabbit). This means that there must be net Ca 2 þ influx between contractions in
rat ventricle, and the [Ca 2 þ ] o trace in Fig. 5 B is actually going below the bath by the
end of the trace to restore the steady state balance before the next beat (
1.5 s
later). Note that during a steady state heartbeat, total Ca 2 þ influx must equal total
Ca 2 þ e
ux (i.e., there is no net gain or loss of Ca 2 þ at the steady state).
Extracellular Ca 2 þ -MEs are also useful for assessing nonsteady state Ca 2 þ fluxes
on a longer time scale ( Bers and MacLeod, 1986; MacLeod and Bers, 1987 ).
Figure 7 A shows that when 0.5 Hz stimulation is stopped there is a very slow
small rise in [Ca 2 þ ] o over many seconds (net Ca 2 þ e
Z
ux), and upon resumption of
stimulation (now at 1 Hz) that there is a net [Ca 2 þ ] o depletion which develops over
Z
A
B
30
30
Rat ventricle
Rabbit ventricle
aNa i = 7.2 mM
aNa i = 7.2 mM
0
30
E Na/Ca
E Na/Ca
E m
60
60
E m
90
Ca efflux
30
0
30
Ca influx
60
0
250
500
0
250
500
Time (ms)
Time (ms)
Fig. 6 Changes in the reversal potential of the Na þ /Ca 2 þ exchange (E Na/Ca ) during the action
potential (E m ) and Ca 2 þ transient in rabbit and rat ventricle. Changes in electrochemical driving force
for Na þ /Ca 2 þ exchange (E Na/Ca
E m ) are shown in the bottom panels, assuming a 3:1 stoichiometry of
Na þ /Ca 2 þ exchanger and aNa i are measured Na þ activity values ( Shattock & Bers, 1989 ). Ca 2 þ
transients driving the contraction are assumed to be the same for both species (resting [Ca 2 þ ] i ¼ 150 nM,
peak [Ca 2 þ ] i ¼ 1 m M, 40 ms after the AP initiation). Note that Ca 2 þ e Z ux is low during rest in
rabbit myocytes because of the low [Ca 2 þ ] i (despite a significant driving force). Based on data in
Shattock & Bers, (1989) , as modified in Bers (2001), with permission.
 
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