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Phylogenetic analyses indicate that SBP / SPL genes with miR156 target
sites can be grouped into five clades ( Guo et al., 2008; Salinas et al., 2012 ).
The phenotypes of plants transformed with miR156-resistant forms of
these genes, as well as their loss-of-function phenotypes, suggest that they
regulate many different aspects of plant development. These include glume
architecture ( Wang et al., 2005 ), inflorescence branching/bract develop-
ment ( Chuck, Whipple, Jackson, & Hake, 2010 ), and ligule differentiation
( Moreno, Harper, Krueger, Dellaporta, & Freeling, 1997 ) in maize; panicle
( Jiao et al., 2010; Miura et al., 2010 ) and grain morphology ( Wang et al.,
2012 ) in rice; fruit ripening in tomato ( Manning et al., 2006 ); and leaf
initiation ( Martin et al., 2010; Wang et al., 2008 ), embryo development
( Nodine & Bartel, 2010 ), pollen development ( Xing, Salinas, Hohmann,
Berndtgen, & Huijser, 2010 ), and trichome ( Gan, Xia, Chen, & Wang,
2011 ) and anthocyanin patterning ( Gou, Felippes, Liu, Weigel, & Wang,
2011 ) in the inflorescence in Arabidopsis . Thus, SBP/SPL genes regulate a
variety of processes
in addition to those associated with vegetative
phase change.
The divergent functions of SPL genes are apparent from the phenotypes
of plants overexpressing these genes ( Cardon et al., 1997; Martin et al., 2010;
Usami et al., 2009; Wang et al., 2008; Wu et al., 2009; Wu & Poethig,
2006 ). However, their normal functions have been difficult to establish
because of the high degree of functional redundancy within this family.
SPL8 ( Unte et al., 2003; Zhang, Schwarz, Saedler, & Huijser, 2007 ),
SPL14 ( Stone, Liang, Nekl, & Stiers, 2005 ), and SPL9 are the only members
of this family in Arabidopsis that have a loss-of-function phenotype on their
own, and of these only SPL9 is regulated by miR156. SPL9 and SPL15 are
members of the same clade. spl9 mutations produce a slight increase in the
rate of leaf initiation and a very weak juvenilized phenotype, and spl15
mutations are nearly wild type in appearance. However, plants mutant for
both genes have a prolonged juvenile phase, are late flowering, have an
increased number of branches, and an increased rate of leaf initiation
( Schwarz, Grande, Bujdoso, Saedler, & Huijser, 2008; Wang et al.,
2008 ). spl9 and spl15 also enhance the male sterility phenotype of spl8 , dem-
onstrating that they act redundantly with this gene to promote pollen devel-
opment ( Xing et al., 2010 ). SPL2 , SPL10 , and SPL11 are also members of a
single clade in Arabidopsis . spl2 mutations have no apparent loss-of-function
phenotype, but slightly enhance the delayed phase change phenotype of the
spl9 spl15 double mutant ( Schwarz et al., 2008 ) and the male sterile pheno-
type of the spl9 spl15 spl8 triple mutant ( Xing et al., 2010 ). SPL10 and
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