The upper side of the angiosperm leaf is specialized for efficient capture
of sunlight whereas the lower side is specialized for gas exchange. In Arab
idopsis, the establishment of polarity in the leaf probably requires the ge
neration and perception of positional information along the radial (adaxial
versus abaxial or central versus peripheral) dimension of the plant. This
is because the future upper (adaxial) side of the leaf develops from cells
closer to the centre of the shoot, whereas the future under (abaxial) side
develops from cells located more peripherally. Here we implicate the Arabid
opsis PHABULOSA and PHAVOLUTA genes in the perception of radial positional
information in the leaf primordium. Dominant phabulosa (phb)(1) and phavolu
ta (phv) mutations cause a dramatic transformation of abaxial leaf fates in
to adaxial leaf fates. They do so by altering the predicted sterol/lipid-bi
nding domains of ATHB14 and ATHB9, proteins of previously unknown function
that also contain DNA-binding motifs. This change probably renders the prot
ein constitutively active, implicating this domain as a central regulator o
f protein function and the PHB and PHV proteins as receptors for an adaxial
izing signal.