Journal of Electrocardiology Vol. 31 Supplement
Cellular Mechanisms for the Slow Phase
of the Frank-Starling Response
Wolfgang F. Bluhm, PhD,* Derrick Sung, MS,~-
Wilbur Y. W. Lew, MD,*$ Alan Garfinkel, PhD,§
and Andrew D. McCulloch, PhD~-
Abstract: Following a step increase in sarcomere length, isometric cardiac muscle
tension increases instantaneously by the Frank-Staffing mechanism. In isolated
papillary muscle and myocytes, there is an additional significant rise in developed
tension over the following 15 min due to an unknown mechanism. This slow
change in tension could not be explained by mechanical heterogeneity of the
muscle preparations or by an increase in myofilament sensitivity to Ca2+. The
slow change in tension was not dependent on sarcoplasmic reticulum Ca2+
loading assessed with rapid cooling contractures, and was not significantly altered
by sarcoplasmic reticulum Ca2+ depletion (ryanodine) or inhibition of sarcoplas-
mic reticulum Ca2+ reuptake (cydopiazonic acid). We used the Luo-Rudy ionic
model of the ventricular myocyte together with a model of the length-dependent
myofilament activation by Cax+ to examine the effects of step changes in the
parameters of sarcolemmal ion fluxes as possible mechanisms for the slow change
in stress. The slow increase in tension was simulated by step changes in the
Na+-K + pump or Na + leak currents, suggesting that the slow change in stress may
be caused by length induced changes in Na + fluxes. The model also predicted a
slow increase in the magnitude of the initial repolarization during phase 1 of the
action potential. The combination of experimental and computational models
used in this investigation represents a valuable technique in eluddating the cellular
mechanisms of fundamental processes in cardiac exaltation-contraction coupling.
Key words: heart, Frank-Starling mechanism, excitation-contraction cou-
pling, Luo-Rudy model.
Active stress in cardiac muscle increases with mus-
de length. This fundamental principle has been
From the *Department of Medicine, University of California, San
Diego, California; t-Department of Bioengineering, University of Cali-
fornia, San Diego, California; ¢Cardiology Division, Department of
Veterans Affairs Medical Center, San Diego, California; and ~Depart-
ments of Physiological Science and Medicine (Cardiology), University of
California, Los Angeles, California.
Supported by NIH grant I-IL-41603, NSF grant BES-9634974,
the Department of Veterans Affairs, and the American Heart
Association.
Reprint requests: Andrew D. McCulloch, PhD, Department of
Bioengineering, Institute for Biomedical Engineering, 9500 Gil-
man Drive, University of California, San Diego, La Jolla, CA
92093-0412.
Copyright © 1998 by Churchill Livingstone ®
0022-0736/98/310S-100355.00/0
known since the classic studies of Frank and Starling
almost a century ago. Over the last two decades, it has
become evident that this increase is primarily caused
by length-dependent activation of the thin filament.
The length-tension relationship has long been consid-
ered a fundamental property of isolated cardiac mus-
cle preparations and has been the subject of numer-
ous review articles (i-5). However, the contractile
force of cardiac musde does not depend on the
instantaneous muscle length alone. Active and pas-
sive forces in cardiac muscle both depend on the time
history of sarcomere length.
In 1973, Parmley and Chuck (6) first showed that
the immediate increase in cardiac muscle tension
13