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Growth, Differentiation and Sexuality

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108 J. Wendl<strong>and</strong> <strong>and</strong> A. Walther<br />

Table. 6.1. (continued)<br />

A. gossypii N. crassa S. cerevisiae<br />

Mitotic exit<br />

AFR035W (0.0) NCU03242.1 (e-053),<br />

NCU07296.1 (e-146)<br />

YNL161W CBK1<br />

ACL006W (0.0) NCU09258.1 (e-063) YMR001C CDC5<br />

ADR033W (0.0) NCU09071.1 (e-135) YGR092W, YPR111W DBF2, DBF20<br />

ADL149W (e-170) NCU00622.1 (e-056) YHR158C, YGR238C KEL1, KEL2<br />

ACR292W (0.0) NCU03379.1 (e-041) YAL024C LTE1<br />

ADR317C (e-175) NCU00978.1 (e-072) YDL028C MPS1<br />

AER425W (e-007),<br />

syntenic homolog<br />

None YNL078W NIS1<br />

Actin ring formation <strong>and</strong> cytokinesis<br />

AFR669W (0.0) NCU01431.1 (e-085) YNL271C BNI1<br />

ABL200W (e-062) NCU00064.1 (e-014) YNL233W BNI4<br />

AFR027C (e-031) None YNL166C BNI5<br />

AFR301C (e-118) None YIL159W BNR1<br />

ABL153W (0.0) NCU04251.1 (0.0) YNL192W CHS1<br />

AEL190W (0.0) NCU05239.1 (0.0) YBR038W CHS2<br />

AEL189W (0.0) NCU09324.1 (0.0),<br />

YBR023C CHS3<br />

NCU04350.1 (e-122)<br />

ACR227W (e-156) NCU02351.1 (e-032),<br />

NCU02592.1 (e-042),<br />

NCU09322.1 (e-076)<br />

YBL061C SKT5/CHS4<br />

ADL288C (0.0) NCU04095.1 (e-045) YDL117W CYK3<br />

ABR082W (e-080) NCU04763.1 (e-017) YMR032W HOF1/CYK2<br />

AFL150C (0.0) NCU03116.1 (e-048) YPL242C IQG1/CYK1<br />

AFL030C (e-051) NCU06617.1 (e-024) YGL106W MLC1<br />

AEL280W (e-022) None YPR188C MLC2<br />

ACR068W (0.0) NCU00551.1 (0.0) YHR023W MYO1<br />

a Ashbya gossypii sequences correspond to the systematic nomenclature (http://agd.unibas.ch/); Neurospora crassa sequences<br />

correspond to the genome annotation available at http://www.broad.mit.edu/annotation/fungi/neurospora_crassa<br />

_7/index.html. Values in brackets indicate the e-values generated by using BLAST-P of the fungal sequences against the S.<br />

cerevisiae protein. An e-value of 0.0 indicates very high sequence identity. Syntenic homology (conserved chromosomal<br />

gene order of homologs) between Ashbya <strong>and</strong> S. cerevisiae genes was used to indicate homology, despite low e-value blast<br />

scores. E-values from Ashbya to S. cerevisiae comparisons were found to be higher than those of N. crassa to S. cerevisiae<br />

comparisons, reflecting the closer relationship of the former species pair<br />

Bud8p <strong>and</strong> Bud5p (Roemer et al. 1996; Kang et al.<br />

2001, 2004a).<br />

B. Placement of the Site of Cell Division<br />

in Schizosaccharomyces pombe<br />

Thecleavageplane,<strong>and</strong>thustheseptumposition<br />

in S. pombe is selected in a manner similar to that<br />

of animal cells, which has made S. pombe amodel<br />

organism to study cell division. However, in animal<br />

cells, selection of the cleavage plane is dependent on<br />

spindle microtubules forming a structure termed<br />

‘midbody’, whereas in S. pombe the mitotic spindle<br />

can be disrupted without interfering with the<br />

selection of the septal site (Chang et al. 1996). This<br />

may reflect a difference in nuclear division, which<br />

occurs as a closed mitosis in S. pombe, in which the<br />

nuclear envelope is not broken down (Chang 2001).<br />

In S. pombe, positioning of the nucleus is important<br />

for the placement of the division site, since<br />

mispositioning of the nucleus is followed by an<br />

aberrant localization of the division plane (Chang<br />

<strong>and</strong> Nurse 1996; Chang et al. 1996; Tran et al. 2001).<br />

TheMid1/Dmf1proteinwasshowntoplayanessential<br />

role in the positioning of the cleavage plane,<br />

<strong>and</strong> may encode a l<strong>and</strong>mark protein defining the<br />

septal site (Sohrmann et al. 1998). Mutants of mid1<br />

are still able to form actin rings, but the positioning<br />

of the ring is r<strong>and</strong>omized at different angles. Mid1p<br />

is localized in the nucleus in interphase cells but exits<br />

the nucleus upon phosphorylation during mitosis,<br />

regulated via the polo-like kinase Plo1 (Ohkura<br />

et al. 1995; Bähler et al. 1998; Fig. 6.1B). Mid1p<br />

then forms a b<strong>and</strong> on the cell cortex in the vicinity<br />

of the nucleus, <strong>and</strong> this localization is dependent<br />

on the position of the nucleus, since nuclear po-

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