The mechanism of var2 leaf variegation

Yu, Fei
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The Arabidopsis f&barbelow;ilamentation t&barbelow;emperature s&barbelow;ensitive (FtsH) metalloprotease gene family comprises 12 members (AtFtH1-AtFtsH12 ), including three pairs of closely related genes that are targeted to chloroplasts (AtFtsH2 and AtFtsH8; AtFtsH1 and AtFtsH5; and AtFtsH7 and AtFtsH9). Mutations in AtFtsH5 ( var1) and AtFtsH2 (var2) give rise to variegated plants with green- and white-sectored leaves. Cells in the green sectors contain morphologically normal chloroplasts, whereas cells in the white sectors are blocked in chloroplast biogenesis. A major question is how chloroplasts arise in cells that have a mutant genotype. We have found by two-dimensional (2-D) green gel and gel filtration analyses that AtFtsH2/VAR2 forms oligomeric complexes. Two bands in the 2-D green gels that correspond to AtFtsH5/VAR1 + AtFtsH1 and AtFtsH2/VAR2 + AtFtsH8 have been identified, and these bands are coordinately reduced in amount in var1 and var2 thylakoids that lack AtFtsH5/VAR1 and AtFtsH2/VAR2, respectively. These reductions are not because of alterations in transcript abunDance; Overexpression of AtFtsH8 in var2-4 (a putative null allele) normalizes the variegation phenotype of the mutant and restores the two bands to their wild-type levels. These results suggest that AtFtsH8 is interchangeable with AtFtsH2/VAR2 in AtFtsH-containing oligomers, and that the two proteins have redundant functions. Similarly, overexpression of AtFtsH1 in var1-1 also restores the variegation to green, indicating that AtFtsH1 and AtFtsH5/VAR1 also share redundant functions. Based on our findings, we propose that AtFtsH1, AtFtsH2/VAR2, AtFtsH5/VAR1 and AtFtsH8 interact to form oligomeric structures, and that subunit stoichiometry is controlled post-transcriptionally in var1 and var2, perhaps by turnover. A threshold model is presented to explain the pattern of variegation in both var2 and var1 in which AtFtsH8 and AtFtsH1 provide compensating activities in the green sectors of the mutant, respectively. To better understand the regulation of leaf variegation and chloroplast development, we carried out a large-scale activation-tagging screen to identify potential var2 modifiers. Results from the characterization of three putative var2 modifiers are presented.

Genetics, development and cell biology, Plant physiology