ELSEVIER Biochimica el BiophysicaActa 1185 (1994) 327-335 ~ ~ ~
Effects of the creatine analogue g-guanidinopropiomc acid
on skeletal muscles of mice deficient in muscle creatine kinase
Jan van Deursen a Paul Jap a, Arend Heerschap b, Henk ter Laak c, Wim Ruitenbeek d,
136 Wieringa a'*
a Depart~wnts of Cell Biology and Histology, b Diagnostic Radiolo~, ~ Neuroto~, and d Paediatrics, Facul O" o f Medical Sciences.
Uniz'ersio' of Nijmegen, P.O. Box 9101, 6500 lib Nijmegen, ITw Netherlands
(Received 4 October 1993)
Abstract
To evaluate the effects of phosphocreatine (PCr) and creatine (Cr) depletion on skeletal muscles of mice deficient in muscle
crcatine kinase (M-CK), we have fed mutant mice a diet containing the creatine analogue ~-guanidinopropionic acid (g3GPA).
After 8-10 weeks of feeding, accumulation of the creatine analogue in M-CK-deficient muscles was comparable to that observed
in muscles of wild-type mice. Strikingly, and unlike wild types, mutants did not accumulate pbosphorylated ~GPA, indicating
that MM-CK is the only muscle CK isoform which can phosphorylate 8GPA. in M-CK-defk-ient muscles there was respective
depletion of PCr, Cr and ATP levels to 31, 4l and 83% of normal. ~ avcra~',. ,-~'~s-sectional area of type 213 fibres in
gastrocnemius muscles was very much reduced and was similar to type I and type 2A fibres which maintained their normal size.
The maximal isometric twitch force developed by gastrocnemius-plantaris-soleus (GPS) muscle complexes of tgGPA-treated
mutants was reduced by about 30%, but these muscles showed an increased fatigue resistance during I and 5 Hz contraction.
Mitoehondrial enzyme activities in the upper hind limb musculature of null mutants were 20-35% increased by the g3GPA diet.
Altogether, these results provide evidence that certain functions of the creatine kinase/phosphocreatine (CK/PCr) system are
not eliminated solely by the loss of M-CK.
Key words: CK/PCr system; Creatine kinase deficiency; Phos,ahoc:c., :,,=: 8-Guanidinopropionic acid; Muscle; Skeletal muscle
bioenergetics; (M-CK-deficient mice)
1. Introduction
Skeletal muscles contain high levels of phospho-
creatine (PCr) and creatine kinase (CK), which are
presumed to function in energy buffering and energy
transport [1-3]. Thus far, two kinds of animal model
have been used to evaluate the physiological impor-
tance of the CK/PCr system in energy metabolism of
skeletal muscle. One model involves the feeding of
synthetic analogues of Cr, such as /3-gnanidinopro-
pionie acid, which competitively inhibit uptake of crea-
fine (Cr) in skeletal muscle cells [4,5]. Rats put on a
* Corresponding author,
0005-2728/oA/$07.00 © 1994 Elsevier Science B.V. All rights reserved
SgDI 0005-2728(94)00016-X
diet containing !%/3GPA (w/w) for 6-8 weeks accu-
mulate high concentrations of phosphorylated g~GPA
(/3GPAP) in their skeletal muscles, and display PCr
and ATP levels which are respectively 12- and 2-fold
-educed compared to normal. A reduction in the flux
of high-energy phosphates between PCr and ATP par-
allels the decline of PCr, as shown by steady-state
nuclear magnetic resonance (NMR) magnetization
transfer experiments [6,7]. The phosphorylated g3GPA
cannot effectively substitute for PCr, because it is a
poor substrate for CK [8]. g3GPA feeding results in a
number of adap*ivc changes in skeletal muscle. Glyco-
gen levels arc elevatt~, while glycolytic enzymes and
glycogen consumption during muscle exercise are re-
duced [7,9]. The aerobic energy-generating capacity of
type 2 fibres is enhanced [9]. Mitochondria in perinu-
clear and subsarcolemmal areas of type 1 fibres fre-
quently contain paracrystailine inclusions and often are