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2.2 Expression of Actin-Encoding Genes<br />

18 Mycorrhizal Development and Cytoskeleton 297<br />

Arabidopsis is known to have ten actin genes, two of which are pseudogenes<br />

(Meagher et al. 1999). Out of the eight expressed actin genes ACT2, ACT8,and<br />

ACT7 are named Arabidopsis vegetative actin genes due to the accumulation<br />

pattern of transcripts. ACT2 is expressed in young and old vegetative tissue,<br />

but ACT8 only in a subset of the organs and tissues expressing ACT2 (An et al.<br />

1996a). ACT7 is expressed in young expanding vegetative tissues and is also<br />

involved in phytohormone responses (Kandasamy et al. 2001). The rest of the<br />

Arabidopsis actin genes appear to be associated with reproductive processes<br />

(An et al. 1996b; Huang et al. 1997; Meagher et al. 1999). In spite of the high<br />

number of actin genes in most <strong>plant</strong> species and the important cell biological<br />

functions of the actin cytoskeleton, only a few analyses about the expression<br />

of different actin genes in root tissue have been performed (McLean et al.<br />

1990) and the effect of mycorrhiza on the expression of actin genes at mRNA<br />

level has not been investigated.<br />

One-dimensional analyses of actin expression at the protein level have<br />

been performed in Pinus ectomycorrhiza. The similar mobility of <strong>plant</strong> and<br />

fungal actins in the immunoblots of 1-D gels from P. sylvestris- S. bovinus<br />

ectomycorrhiza made it difficult to record the contribution of each symbiotic<br />

partner to the actin signal (Timonen et al. 1993). The presence of the actin signal<br />

throughout the different developmental stages of ectomycorrhiza was suggested<br />

to be a reliable marker for still metabolically active symbiosis (Timonen<br />

et al. 1996).<br />

In the immunoblots of 2-D gels four actin polypeptides were detected in P.<br />

sylvestris radicles, main roots and first order laterals and in short roots. The<br />

polypeptides were also detected in P. sylvestris- S. bovinus young and dichotomous<br />

mycorrhizal short roots. In fully mature coralloid mycorrhiza only two<br />

actin polypeptides occurred. Whether they represented <strong>plant</strong> or fungal actin<br />

or both was not possible to conclude. The presence of four actin polypeptides<br />

in Pinus root tissues is in agreement with the occurrence of a similar number<br />

of actin polypeptides in Vicia faba roots (Janssen et al. 1996), while in the<br />

roots of Phaseolus vulgaris one and two actin polypeptides were detected in<br />

symbiotic root nodules and in uninfected roots, respectively (Pérez et al.<br />

1994).<br />

In the immunoblots of 2-D gels of the vegetative hyphae of S. bovinus two<br />

actin polypeptides occurred that were also detected in ectomycorrhiza (Niini<br />

et al. 1996). Recently, two actin-encoding genes, Sbact1 and Sbact2,were isolated<br />

from S. bovinus nonsymbiotic hyphae (Tarkka et al. 2000). Northern<br />

hybridization with specific probes for each actin gene of S. bovinus indicated<br />

that both actins are expressed in vegetative hyphae and ectomycorrhiza. In<br />

vegetative hyphae, the expression rate and protein level of Sbact1 was tenfold<br />

higher than that of Sbact2. A ten times higher accumulation of Sbact1 than<br />

Sbact2 mRNA was also observed in ectomycorrhizal hyphae, although the

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