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308<br />

Marjatta Raudaskoski, Mika Tarkka and Sara Niini<br />

solin-like headpiece, villin may also act as an actin-severing protein, although<br />

this activity has not yet been demonstrated for <strong>plant</strong> villins.<br />

In Arabidopsis, 17 myosin encoding genes have been identified, which on<br />

the basis of phylogenetic analysis, fall into <strong>plant</strong>-specific myosin classes VIII<br />

(4 genes) and XI (13 genes). The three cloned myosins from maize and four<br />

from Helianthus annuus also fall into these classes (Reddy and Day 2001). The<br />

true structure, enzymatic properties, intracellular localization and physiology<br />

of <strong>plant</strong> myosin are not yet known.<br />

The structure and function of actin binding proteins in association with<br />

the reorganization of the actin cytoskeleton in <strong>plant</strong> cells at mycorrhiza formation<br />

has not yet been studied, but seems to require attention. In endomycorrhiza,<br />

rearrangement of actin cytoskeleton was observed at the colonization<br />

of tobacco cortical cells by endomycorrhizal fungus (Genre and Bonfante<br />

1997). Actin cables, typical for noncolonized cortical root cells, disappeared<br />

and MF polymerized to a network at the plasma membrane surrounding the<br />

arbuscule branches. The observed changes may be hypothesized to require<br />

both the activation and function of proteins involved in the reorganization of<br />

the actin cytoskeleton as a consequence of the perception of signals from the<br />

intruding fungus.<br />

The research on actin binding proteins in filamentous fungi seems to be<br />

restricted to myosin (McGoldrick et al. 1995), in Aspergillus nidulans and to<br />

actin-related proteins Arp1 (Plamann et al. 1994) in Neurospora crassa,<br />

although in yeast, Saccharomyces cerevisiae, most actin binding proteins previously<br />

described in <strong>plant</strong>s are present and have been studied in detail<br />

(Ayscough 1998). In A. nidulans, the myosin I encoding gene myoA has been<br />

cloned and different mutational studies have indicated that MYOA is necessary<br />

for the maintenance of polarized growth, secretion and endocytosis<br />

(McGoldrick et al. 1995; Osherov et al. 1998; Yamashita and May 1998). The<br />

yeast S. cerevisiae and Schizosaccharomyces pombe, similar to animal cells,<br />

contain myosins from classes II and V in addition to those belonging to class<br />

I which predicts that additional myosins will be detected in future from filamentous<br />

fungi.<br />

7 Microtubule-Associated Proteins<br />

7.1 Plant Cells<br />

Microtubule-associated proteins (MAPs) are divided into structural and<br />

motor proteins. Several genes encoding structural <strong>plant</strong> MAPs have been isolated<br />

in the last few years. One of the MAP encoding genes was isolated from<br />

Arabidopsis, where its heat-sensitive mutation resulted in the disintegration of<br />

cortical microtubules in leaf, hypocotyl and root cells. The gene was named

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