Agriculture Reference
In-Depth Information
Table 6.1
Genes involved in
Arabidopsis
meristem formation and maintenance
BELLRINGER
(
BLR
)/
PENNYWISE
(
PNY
)
The
blr
/
pny
mutant phenotype is similar to the
brevipedicellus
mutant
(see text) having both short internodes resulting in a stunted appearance
and excessive development of axillary meristems (Byrne
et al.
, 2003;
Smith & Hake, 2003). Although
blr
/
pny
mutants have normal SAMs, in
different genetic backgrounds
BLR
/
PNY
is required for meristem
maintenance. For instance
blr
/
pny
enhances the weak
stm
mutant
phenotype and is required for meristem maintenance in an
as1 stm
mutant
background (Bryne
et al.
, 2003). BLR/PNY encodes a
BELL1
-like
homeodomain transcription factor physically interacts with the STM and
KNAT1 proteins, resulting in high affinity binding to target sequences
(Byrne
et al.
, 2003; Smith & Hake, 2003).
CUP-SHAPED
COTYLEDON1
(
CUC1
)
CUP-SHAPED
COTYLEDON2
(
CUC2
)
CUP-SHAPED
COTYLEDON3
(
CUC3
)
CUC1
,
CUC2
and
CUC3
encode plant-specific NAC domain proteins with
extensive homology to the
NO APICAL MERISTEM
(
NAM
) gene of
Petunia
and the recently identified
CUPILLIFORMIS
(
CUP
) gene of
Antirrhinum
(Souer
et al.
, 1996; Aida
et al.
, 1997; Takada
et al.
, 2001;
Vroemen
et al.
, 2003; Weir
et al.
, 2004). All three genes are required
redundantly to promote the formation of the shoot apical meristem and the
separation of the cotyledons during embryogenesis. In the developing
embryo all three
CUC
genes are expressed in cells that will form the SAM
and are then subsequently expressed at the boundary between organ
primordia and the meristem during post-embryonic development. Based
on the expression of these genes during embryogenesis and their
combined mutant phenotype, it has been proposed that they either
promote
STM
expression directly or provide signals that establish a
programme of SAM development (Aida
et al.
, 1997, 1999; Long &
Barton, 1998; Takada
et al.
, 2001; Vroemen
et al.
, 2003). Consistent
with this is the formation of ectopic meristems on cotyledons of plants
expressing
CUC1
ubiquitiously (Takada
et al.
, 2001).
DORNROSCHEN
(
DRN
)/
ENHANCER
OF SHOOT
REGENERATION1
(
ESR1
)
Identified as a gain-of-function mutation,
DRN
/
ESR1
is expressed in lateral
organs and the meristem and encodes an AP2/EREB transcription factor.
Misexpression in cultured roots leads to accelerated regeneration of
shoots (Banno
et al.
, 2001), whereas increased
DRN
/
ESR1
levels in shoots
causes an enlargement of the meristem and the successive radialisation of
leaves, before the arrest of the meristem. DRN/ESR1 regulates
STM
,
CLV3
and
WUS
expression and functions to repress stem cell fate. The
role of DRN/ESR1 in organ formation is unclear, but appears to be
required non-cell autonomously (Kirch
et al.
, 2003).
FASCIATA1
(
FAS1
)
FASCIATA2
(
FAS2
)
Mutations in these genes severely disrupt cellular organisation of the SAM
and RAM, leading to fasciation (Leyser & Furner, 1992). In the SAM,
WUS
expression expands both laterally and apically into all three layers.
FAS1/2 encode subunits of the
Arabidopsis chromatin assembly factor-1
(CAF-1) and presumably functions in maintaining epigenetic states during
cell division within meristems (Kaya
et al.
, 2001).
FOREVER YOUNG
(
FEY
)
The
fey
mutant has an altered phyllotaxis, and produces radially symmetric
organs at a high frequency. In addition cells within the vegetative
meristem and leaf primordia are more vacuolated than normal.
FEY
encodes a nodulin-like protein, although how this protein functions in
meristem and leaf development has yet to be determined (Callos
et al.
,
1994)