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Auxin


Introduction
jifri_introPlant development is characterised by a remarkable adaptability to different environmental conditions. Plants have the ability for extensive post-embryonic development involving the activity of stable stem cell populations (meristems), de novo organ formation and changes in growth direction. This provides exceptional flexibility in terms of growth and survival. Many of these developmental events are characterized by an asymmetric distribution (gradient) of the plant signalling molecule auxin. Auxin is unique among plant signalling molecules, because it is transported in a polar fashion through plant tissues. This transport is driven by the action of influx and efflux carrier proteins represented by the AUX1/LAX and PIN protein families, respectively. The subcellular localisation of PIN efflux carriers determines the direction of auxin flow, a mechanism, which can also define asymmetric auxin distribution within tissues. The auxin transport system thus provides positional and directional information for many aspects of adaptive plant development. It also represents a unique model system to study the functional link between basic cellular processes, such as vesicle trafficking or cell polarity establishment, and their developmental outcome at the level of the multicellular plant organism.
 
Polar auxin transport
1mechanismPolar auxin transport is crucial in regulating plant development as it generates asymmetric auxin distributions (auxin gradients) that underlies many different developmental processes. Classical models postulate that auxin movement between cells requires auxin influx and efflux carriers.
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Polar targeting in plants
2polarityThe key feature of polar auxin transport – its controlled directionality - was postulated to result from the asymmetric subcellular localisation of the efflux carriers. Remarkably, as predicted, PIN proteins display an asymmetric localisation within auxin transport competent cells and this polar localisation correlates with the known directions of auxin flow (Fig. 3). Indeed, polarity of PIN proteins has been shown to determine the direction of auxin flow (Wisniewska et al., 2006).
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Endocytosis and recycling in plants
3endocyticrecyclingPolar targeting of PIN proteins is related to their continuous subcellular movement between endosomes and the plasma membrane (Geldner et al., 2001).

PIN internalization occurs by the clathrin-dependent endocytosis mechanism (Dhonukshe et al., 2007) and its recycling back to the plasma membrane requires ARF GEF regulators of vesicle trafficking (Geldner et al., 2001). The constitutive cycling of auxin transport components provides an entry point for internal and external signals, which in this manner can rapidly modulate PIN polarity.

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