Specification of bundle sheath cell fates during maize leaf development: roles of lineage and positional information evaluated through analysis of the tangled1 mutant.

In leaves of the maize tangled1 (tan1) mutant, clusters of bundle sheath (BS)-like cells extend several cells distant from the veins, in association with the single layer of BS cells around the vein. We show that the BS-like cell clusters in tan1 leaves result from the continued division of cells in the procambial/BS cell lineage that do not divide further in wild-type leaves. The ectopic BS-like cells accumulate the BS marker NADP-dependent malic enzyme but not the mesophyll cell marker phosphoenolpyruvate carboxylase, and exhibit thickened walls, suggesting that they differentiate as C4-type BS cells. We propose that bundle sheath cell fate can be conferred on some derivatives of procambial cell divisions in a manner that is heritable through multiple cell divisions and is position-independent.