Elizabeth
 A. Kellogg, Paulo E. A. S. Camara, Paula J. Rudall, Philip Ladd, Simon 
T. Malcomber, Clinton J. Whipple and Andrew N. Doust 
Abstract
The shoot apical meristem of grasses produces the primary branches of 
the inflorescence, controlling inflorescence architecture and hence seed
 production. Whereas leaves are produced in a distichous pattern, with 
the primordia separated from each other by an angle of 180°, 
inflorescence branches are produced in a spiral in most species. The 
morphology and developmental genetics of the shift in phyllotaxis have 
been studied extensively in maize and rice. However, in wheat, Brachypodium,
 and oats, all in the grass subfamily Pooideae, the change in 
phyllotaxis does not occur; primary inflorescence branches are produced 
distichously. It is unknown whether the distichous inflorescence 
originated at the base of Pooideae, or whether it appeared several times
 independently. In this study, we show that Brachyelytrum, the 
genus sister to all other Pooideae has spiral phyllotaxis in the 
inflorescence, but that in the remaining 3000+ species of Pooideae, the 
phyllotaxis is two-ranked. These two-ranked inflorescences are not 
perfectly symmetrical, and have a clear “front” and “back;” this 
developmental axis has never been described in the literature and it is 
unclear what establishes its polarity. Strictly distichous 
inflorescences appear somewhat later in the evolution of the subfamily. 
Two-ranked inflorescences also appear in a few grass outgroups and 
sporadically elsewhere in the family, but unlike in Pooideae do not 
generally correlate with a major radiation of species. After production 
of branches, the inflorescence meristem may be converted to a spikelet 
meristem or may simply abort; this developmental decision appears to be 
independent of the branching pattern. 
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