The active transport of the plant hormone auxin plays a major role in the initiation of organs at the shoot apex. Polar localized membrane proteins of the PIN1 family facilitate this transport, and recent observations suggest that auxin maxima created by these proteins are at the basis of organ initiation. This hypothesis is based on the visual, qualitative characterization of the complex distribution patterns of the PIN1 protein in Arabidopsis. To take these analyses further, we investigated the properties of the patterns using computational modeling. The simulations reveal previously undescribed properties of PIN1 distribution. In particular, they suggest an important role for the meristem summit in the distribution of auxin. We confirm these predictions by further experimentation and propose a detailed model for the dynamics of auxin fluxes at the shoot apex.auxin Í modeling Í shoot meristem T here is strong evidence that active auxin transport, generated by influx and efflux carriers, creates patterns of auxin distribution at the shoot apex. This distribution is, in turn, interpreted in terms of differential growth and cell differentiation (1-3). In Arabidopsis, AUX1, a putative influx transporter (4), is mainly located in the surface layer (L1) of the shoot apical meristem (2) (Fig. 1A). Interestingly, the protein seems to be homogeneously distributed in plasma membranes of the individual cells. Therefore, it has been proposed that AUX1 helps to restrict auxin to these layers, although additional mechanisms may be required (5). The efflux facilitator PIN1 also is localized in the surface layers of the meristem, but in contrast to AUX1 it is often localized on certain anticlinal sides of the cells only. Because neighboring cells often show coherent PIN1 positioning, it was proposed that PIN1 is responsible for directed hormone flows within the meristem L1 layer (Fig. 1 A). In particular, careful immunological studies have revealed that the membranes carrying PIN1 are preferentially oriented toward the incipient primordia, suggesting auxin transport toward the young organs (2, 3).Together, the observations so far suggest a dynamic scenario where auxin is transported to the meristem from basally localized tissues via the L1 layer. At the meristem surface, auxin is redistributed and accumulates at particular sites where it will induce the initiation of new organs. This accumulation subsequently leads to the activation of transport in the provascular tissues causing an inward directed flow (Fig. 1B). The young organ is thus transformed into an auxin sink, which depletes its surroundings from auxin and prevents the formation of new primordia in its vicinity.Although this scenario is relatively straightforward, the previous observations leave a number of questions open. First, it is not clear at all why auxin should start to accumulate at the site where a primordium will be initiated. Second, the immunolabelings reveal a very complex distribution of PIN1 proteins (Fig. 2). As a result, the interpretation of these...