Cortical polarity ensures its own asymmetric inheritance in the stomatal lineage to pattern the leaf surface.

Asymmetric cell divisions specify differential cell fates across kingdoms. In metazoans, preferential inheritance of fate determinants into one daughter cell frequently depends on polarity-cytoskeleton interactions. Despite the prevalence of asymmetric divisions throughout plant development, evidence for analogous mechanisms that segregate fate determinants remains elusive. Here, we describe a mechanism in the Arabidopsis leaf epidermis that ensures unequal inheritance of a fate-enforcing polarity domain. By defining a cortical region depleted of stable microtubules, the polarity domain limits possible division orientations. Accordingly, uncoupling the polarity domain from microtubule organization during mitosis leads to aberrant division planes and accompanying cell identity defects. Our data highlight how a common biological module, coupling polarity to fate segregation through the cytoskeleton, can be reconfigured to accommodate unique features of plant development.

Polarizing to the challenge: New insights into polarity-mediated division orientation in plant development.

Land plants depend on oriented cell divisions that specify cell identities and tissue architecture. As such, the initiation and subsequent growth of plant organs require pathways that integrate diverse systemic signals to inform division orientation. Cell polarity is one solution to this challenge, allowing cells to generate internal asymmetry both spontaneously and in response to extrinsic cues. Here, we provide an update on our understanding of how plasma membrane-associated polarity domains control division orientation in plant cells. These cortical polar domains are flexible protein platforms whose positions, dynamics, and recruited effectors can be modulated by varied signals to control cellular behavior. Several recent reviews have explored the formation and maintenance of polar domains during plant development [1-4], so we focus here on substantial advances in our understanding of polarity-mediated division orientation from the last five years to provide a current snapshot of the field and highlight areas for future exploration.