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glycolytic pathway and LeT6 , which encodes a homeobox transcription factor. The
mutant shows more compound leaves than in wild-type plants. Grafting of wild-type
shoots onto stocks of the dominant mouse ears mutant caused the morphology of
leaves at the apex of the plant to appear similar to those of mouse ears mutants, and
this correlated with the presence of increased abundance of mouse ears mRNA in
the wild-type shoot (Kim et al. , 2001).
Positioning the floral stimulus within a regulatory hierarchy will allow it to be
approached using the molecular genetic tools available in Arabidopsis . Screening
for mutations that suppress the early-flowering phenotype caused by expressing
CO or FT in the phloem, or screening randomly for further proteins and RNAs
that influence flowering-time when expressed in the phloem, is likely to identify
further components of the system. Similarly, use of full genome microarrays will
enable identification of all of the genes whose expression is induced by overexpres-
sion of CO in the phloem, and therefore provide leads as to the identity of mobile
components.
7.3
Intercellular communication during floral development
Dramatic changes in gene expression occur at the apex of the plant on the transition
to flowering. Recently, these changes were analyzed using microarrays containing
oligonucleotides derived from almost all Arabidopsis genes, and within 7 days of
a shift from short to long day lengths, the expression of 332 genes was induced or
repressed at least twofold in both the Landsberg erecta and Columbia accessions
(Schmid et al. , 2003). Perhaps surprisingly, of these 332 genes, the majority (231)
showed reduced expression during floral induction, whereas the remainder were
increased. At least one of the repressed genes, encoding an AP2-like transcription
factor, was shown to actively repress flowering when overexpressed, suggesting
that its downregulation during flowering may play an active role in floral induction
(Schmid et al. , 2003). Similarly, some of the earliest genes that show increased ex-
pression during floral induction appear to promote flowering. For example, the SOC1
gene of Arabidopsis is induced at the meristem within a few hours of shifting plants
from short to long days (Borner et al. , 2000; Samach et al. , 2000), and the orthol-
ogous gene in Sinapis alba behaves similarly (Bonhomme et al. , 2000). Mutations
in SOC1 delay flowering, but do not have an effect on floral development (Borner
et al. , 2000; Lee et al. , 2000; Samach et al. , 2000). In contrast, AP1 and LFY ,two
genes that confer floral meristem identity on the developing floral primoridum, are
among the first genes with established roles in floral development whose transcripts
increase in abundance at the meristem (Mandel et al. , 1992; Weigel et al. , 1992).
Mutations in these genes cause shoots or flowers with shoot characters to develop
from primordia that would normally form flowers, and the proteins encoded by these
genes are involved in activation of downstream transcription factors that confer or-
gan identity on floral organs (Weigel & Meyerowitz, 1993). Recent work indicates
that LFY and orthologous genes in other species, as well as some of the transcription
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