European Journal of Pharmacology, 237 (1993) 83-92 83
© 1993 Elsevier Science Publishers B.V. All rights reserved 0014-2999/93/$06.00
EJP 53121
Influence of Ca 2+ channel agonists and antagonists
on the post-rest staircase
Simon Harrison, Richard Milner and Mark Boyett
Department of Physiology, Unicersityof Leeds, Leeds LS2 9JT, UK
Received 14 December 1992, revised MS received 15 March 1993, accepted 23 March 1993
It is now generally accepted that the post-rest staircase consists of two phases, an initial fast phase occurring over the first five
to six beats followed by a slow phase which occurs over a period of minutes. In this study, the influence of Ca 2+ channel
antagonists (Cd 2+ and nifedipine) and agonists (Bay K 8644 and noradrenaline) on developed tension is investigated to assess
the role played by the L-type Ca 2+ current (ica) during post-rest recovery. The results show that, at a range of stimulus
frequencies, exposure of guinea-pig papillary muscles to either Cd 2+ or nifedipine greatly reduces the fast phase of the staircase
and reduces the slow phase of the staircase compared to control (both effects being more prominent at higher stimulus
frequencies). Exposure of guinea-pig papillary muscles to Bay K 8644 or noradrenaline potentiated both the fast and slow phases
of the staircase with greater proportionate effects observed at higher stimulus frequencies. These results suggest that ica plays an
important role in both phases of the staircase. The first post-rest beat (the rested state contraction) proved very resistant to Ca 2+
channel blockade suggesting that iCa plays little role in this contraction which may be the result of entry of Ca 2+ into the cell via
a relatively Cd 2+ insensitive pathway, for example the Na+-Ca 2+ exchanger.
Ca 2+ currents; Heart; Contraction
I. Introduction
Bowditch (1871) was the first to describe the changes
in contractility of cardiac muscle during stimulation
following a period of rest. He observed that the first
post-rest beat was small but subsequent beats then
increased until a steady state was reached, a phe-
nomenon he called 'Treppe' (lit: staircase). In more
recent years this phenomenon has been investigated to
determine the underlying ionic mechanisms. About 20
years ago two hypotheses were proposed. Langer (1968)
suggested that changes in developed tension (the ten-
sion developed during the twitch contraction) during
post-rest stimulation results from an increase in the
intracellular sodium activity (Na ÷) induced by repeti-
tive activity. An increase in Na ÷ will lead to increased
Ca 2+ loading of the preparation via the Na+-Ca 2+
exchanger. In contrast, Kaufmann et al. (1974), using
an analog computer model based on movements of
Ca 2+ alone, were able to simulate the rapid changes in
developed tension associated with increasing stimulus
Correspondence to: M. Boyett, Department of Physiology, University
of Leeds, Leeds LS2 9JT, UK. Tel. (44) (0532) 334265, fax (44) (0532)
334248.
frequency, suggesting that Ca 2+ loading via the L-type
Ca 2+ current (ica) plays an important role. Further-
more, Lakatta and Spurgeon (1980) have reported that
the staircase is extensively modified by changing extra-
cellular [Ca 2+] and is sensitive to verapamil and iso-
proterenol. These results suggest an important role for
ica in the generation of the staircase.
However, it is now generally accepted that the in-
crease in contraction following a rest can be divided
into two phases (e.g. Harrison et al., 1992). The first
phase is rapid and occurs over the first five to six beats
and this is followed by a slower phase which takes
place over a period of minutes. This suggests that more
than one mechanism is involved in the increase in
developed tension after a period of rest.
Lee (1987) and Hryshko and Bers (1990) measured
ic~ during post-rest stimulation. Their results indicate
that the amplitude of ica increases during the first few
beats following a rest period, a time which coincides
with the fast phase of the staircase. Therefore, it is
tempting to suggest that changes in ic~ , may underlie
the fast phase of the staircase. The model of Kauf-
mann et al. (1974), was able to simulate the fast phase
of the staircase but was not able to simulate slow
changes in 'developed tension'. This is consistent with
ic~ being responsible for the fast phase, but it also