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Dictyostelium
Myosin Heavy Chain Kinases
Myosin II filament assembly in=
Dictyostelium is regulated via Myo=
sin
heavy chain (MHC) phosphorylation through the combined activities of at lea=
st
one MHC phosphatase and a group of MHC kinases (De la Roche et al., 2002; Yumura et al., 2005). Myosin Heavy Chain Kinase A was the
first of the heavy chain kinases to be identifi=
ed and
purified, and cellular and biochemical studies of MHCK A demonstrated that =
the
activity of this kinase drives myosin filament disassembly both in vitro and in vivo by phosphorylating the regu=
latory
threonine sites of MHC (Cote and Bukiejko,
1987). Cloning and sequence analysis of t=
he MHCK
A gene revealed a novel, modular organization to the protein (Figure 3) (Futey et
al., 1995). The amino-terminal domain of MHCK =
A is
predicted to have significant a-helical coiled-coil structure and has been shown biochemically to mediate MHCK A oligomerization (Kolman=
span>
and Egelhoff, 1997). The central catalytic region of MH=
CK A
is the prototype for a novel family of protein kinases=
,
referred to as &q=
uot;a-kinases&q=
uot;,
which do not exhibit any apparent sequence similarity to any of the
conventional protein kinase families (Drennan<=
/span>
and Ryazanov, 2004; Egelho=
ff,
1999; Futey et al., 1995). Representatives of this kinase fam=
ily
have been identified in diverse groups of organisms including several kinases in Di=
ctyostelium
(Yumura=
span> et
al., 2005), as well as others in mammali=
an
systems that include eukaryotic elongation factor-2 ki=
nases
and the Channel-kinase TRPM7/ChaK1 (Drennan<=
/span>
and Ryazanov, 2004). At its carboxyl-terminus, MHCK A
possesses a domain containing a 7-fold WD
repeat motif (Sondek=
span>
and Siderovski, 2001) that facilitates phosphorylat=
ion of
myosin heavy chain by physically targeting MHCK A directly to myosin II
filaments (Steimle et al., 2001a)=
.
Localization studies in chemotaxing cells revealed that MHCK A is recruited to
sites of pseudopod formation at the cell anteri=
or
(Figure 5) (Steimle et al., 2001b)=
. In the context of cytokinesis, MHC=
K A is
enriched at the actin-rich polar regions of the dividing cell (Liang et
al., 2002; Nagasaki et al., 2002). The accumulation of MHCK A at acti=
n-rich
cell extensions, coupled with reduced levels of myosin II at such sites (De la Roche et al., 2002; Wessels et al., 1988), suggests a model in which my=
osin
II filament disassembly is promoted at sites of MHCK A enrichment, and thus=
myosin
II-based cross-linking of actin filaments at regions forming cellular
extensions may be relaxed. In turn, this may promote the assembly and
reorganization of the actin cytoskeleton necessary for the formation of pseudopodial structures (Heid et
al., 2004; Laevsky and Kne=
cht,
2003; Steimle et al., 2001b; Stites et al., 199=
8).
Analyses of the Dictyostelium
genome data base have revealed that there are at least 6 members of the
alpha-kinase family in Dictyosteliu=
m (Kreppel<=
/span>
et al., 2004). Recent studies by Yumura
and colleagues indicate that of the Dictyostelium
alpha kinases, only three - MHCK A, MHCK B (Rico and Egelhoff,
2003), and MHCK C (Liang et
al., 2002; Nagasaki et al., 2002) - appear to have physiologica=
lly
significant functions in regulating myosin II filament disassembly in the c=
ell (Yumura=
span> et
al., 2005). Upon initial examination, MHCK A, =
B, and
C appear to represent functionally redundant activities since they share a
common ability to specifically phosphorylate the
regulatory threonine residues in the tail of the myosin II heavy chain and
drive the disassembly of myosin II bipolar filaments both in vitro and in the cell (Figure 4).
The functional differences amo=
ng the
MHCKs are reflected, in part, by their distinct
localization patterns during dynamic cellular processes such as cell migrat=
ion
and cytokinesis (Figure 5). I=
n a migrating
cell, for example, MHCK A is preferentially enriched at sites of pseudopod formation at the cell anterior where the ki=
nase
is expected to catalyze the disassembly of myosin II-based cross-linking of
actin filaments which, in turn, may facilitate the reorganization of the ac=
tin
cytoskeleton necessary for the formation of cellular extensions (Steimle et al., 2001b)=
.
In contrast, MHCK C co-localizes with myosin II at
the trailing edge of migrating cells, suggesting a role for this enzyme in =
the
coordinated turnover of myosin II filaments that mediate contraction of the=
cell
rear (Liang et
al., 2002; Nagasaki et al., 2002). During cytokinesis, MHCK C exhibits
robust localization to the cleavage furrow. In contrast, MHCK A displays a per=
sistent
localization at the poles of the dividing cell and at the “anterior=
8221;
of the migrating daughter cells (Liang et
al., 2002; Nagasaki et al., 2002). Recent studies indicate that MHCK =
B,
like MHCK C, is recruited to the cleavage furrow during cytokinesis =
(Yumura=
span> et
al., 2005). It is particularly interesting tha=
t MHCK
B and MHCK C localizations depend on the presence of myosin II in the cell =
(Nagasaki et al., 2002; Yumura et al., 2005); this contrasts with MHCK A
localization which does not require myosin II but does depend on the presen=
ce
of filamentous actin in the cell (Steimle et al., 2001b)=
. Taken together, these results sugg=
est
that while the three kinases share similar cata=
lytic
activities, they appear to have distinct functions in regulating myosin II
filament assembly in different cellular contexts.
References
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yosin
heavy chain kinase from Dictyostelium discoideum. J Biol Chem. 262:1065-72.
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T.T. Egelhoff, and G.P. Cote. 2002. Signaling pathways regulating Dictyostel=
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myosin II. J Mu=
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Drennan, D., and A.G. Ryazanov. 2004. Alpha-kinases:
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Egelhoff, T.T., and =
Cote, G.P.
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=
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