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Self-regulatory patterning mechanisms capable of generating biologically meaningful, yet unpredictable cellular patterns offer unique opportunities for obtaining mathematical descriptions of underlying patterning systems properties. The networks of higher-order veins in leaf primordia constitute such a self-regulatory system. During the formation of higher-order veins, vascular precursors are selected from a homogenous field of subepidermal cells in unpredictable positions to eventually connect in complex cellular networks. Auxin transport routes have been implicated in this selection process, but understanding of their role in vascular patterning has been limited by our inability to monitor early auxin transport dynamics  in vivo . Here we describe a live-imaging system in emerging  Arabidopsis thaliana  leaves that uses a  PIN1:GFP  reporter to visualize auxin transport routes and an  Athb8:YFP  reporter as a marker for vascular commitment. Live-imaging revealed common features initiating the formation of all higher-order veins. The formation of broad  PIN1  expression domains is followed by their restriction, leading to sustained, elevated  PIN1  expression in incipient procambial cells files, which then express  Athb8 . Higher-order  PIN1  expression domains (hPEDs) are initiated as freely ending domains that extend toward each other and sometimes fuse with them, creating connected domains. During the restriction and specification phase, cells in wider hPEDs are partitioned into vascular and non-vascular fates: Central cells acquire a coordinated cell axis and express elevated  PIN1  levels as well as the pre-procambial marker  Athb8 , while edge cells downregulate  PIN1  and remain isodiametric. The dynamic nature of the early selection process is underscored by the instability of early hPEDs, which can result in dramatic changes in vascular network architecture prior to  Athb8  expression, which is correlated with the promotion onto vascular cell fate.

Dynamic auxin transport patterns preceding vein formation revealed by live-imaging of Arabidopsis leaf primordia