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Gravity is a major cue for the proper growth and development of plants. The response of plants to gravity implies starch-filled plastids, the statoliths, which sediments at the bottom of the gravisensing cells, the statocytes. Statoliths are assumed to modify the transport of the growth hormone, auxin, by acting on specific auxin transporters, PIN proteins. However, the complete gravitropic signaling pathway from the intracellular signal associated to statoliths to the plant bending is still not well-understood. In this article, we build on recent experimental results showing that statoliths do not act as gravitational force sensor, but as position sensor, to develop a bottom-up theory of plant gravitropism. The main hypothesis of the model is that the presence of statoliths modifies PIN trafficking close to the cell membrane. This basic assumption, coupled with auxin transport and growth in an idealized tissue made of a one-dimensional array of cells, recovers several major features of the gravitropic response of plants. First, the model provides a new interpretation for the response of a plant to a steady stimulus, the so-called sine-law of plant gravitropism. Second, it predicts the existence of a gravity-independent memory process as observed recently in experiments studying the response to transient stimulus. The model suggests that the timescale of this process is associated to PIN turnover, calling for new experimental studies.

Gravity is a major cue for the proper growth and development of plants. The response of plants to gravity implies starch-filled plastids, the statoliths, which sediments at the bottom of the gravisensing cells, the statocytes. Statoliths are assumed to modify the transport of the growth hormone, auxin, by acting on specific auxin transporters, PIN proteins. However, the complete gravitropic signaling pathway from the intracellular signal associated to statoliths to the plant bending is still not well-understood. In this article, we build on recent experimental results showing that statoliths do not act as gravitational force sensor, but as position sensor, to develop a bottom-up theory of plant gravitropism. The main hypothesis of the model is that the presence of statoliths modifies PIN trafficking close to the cell membrane. This basic assumption, coupled with auxin transport and growth in an idealized tissue made of a one-dimensional array of cells, recovers several major features of the gravitropic response of plants. First, the model provides a new interpretation for the response of a plant to a steady stimulus, the so-called sine-law of plant gravitropism. Second, it predicts the existence of a gravity-independent memory process as observed recently in experiments studying the response to transient stimulus. The model suggests that the timescale of this process is associated to PIN turnover, calling for new experimental studies. [ABSTRACT FROM AUTHOR]

Abstract: Cell-to-cell communication is absolutely essential for multicellular organisms. Both animals and plants use chemicals called hormones for intercellular signaling. However, multicellularity of plants and animals has evolved independently, which led to establishment of distinct strategies in order to cope with variations in an ever-changing environment. The phytohormone auxin is crucial to plant development and patterning. PIN auxin efflux carrier-driven polar auxin transport regulates plant development as it controls asymmetric auxin distribution (auxin gradients), which in turn modulates a wide range of developmental processes. Internal and external cues trigger a number of posttranslational PIN auxin carrier modifications that were demonstrated to decisively influence variations in adaptive growth responses. In this review, we highlight recent advances in the analysis of posttranslational modification of PIN auxin efflux carriers, such as phosphorylation and ubiquitylation, and discuss their eminent role in directional vesicle trafficking, PIN protein de-/stabilization and auxin transport activity. We conclude with updated models, in which we attempt to integrate the mechanistic relevance of posttranslational modifications of PIN auxin carriers for the dynamic nature of plant development. [Copyright &y& Elsevier]

The monocot coleoptile tip region has been generally supposed to be the source of IAA to supply IAA to basal parts by the polar IAA transport system, which results in gravi- and phototropic curvature of coleoptiles. Based on this IAA transport system and gravitropism of maize coleoptiles, we have developed two screening methods to identify small molecules from a large chemical library that inhibit IAA transport. The methods detect molecules that affect (i) gravitropic curvature of coleoptiles; and (ii) the amount of IAA transported from the tip. From 10,000 chemicals, eight compounds were identified and categorized into two groups. Four chemicals in group A decreased IAA transport from the tip, and increased endogenous IAA levels in the tip. The structures of two compounds resembled that of 1-N-naphthylphthalamic acid (NPA), but those of the other two differed from structures of known IAA transport inhibitors. Four chemicals in group B strongly inhibited IAA transport from the tip, but IAA levels at the tip were only slightly affected. At higher concentrations, group B compounds inhibited germination of Arabidopsis, similarly to brefeldin A (BFA). Analysis of the cellular distribution of PIN2–green fluorescent protein (GFP) and PIN1–GFP in Arabidopsis revealed that one of the four chemicals in group B induced internalization of PIN1 and PIN2 proteins into vesicles smaller than BFA bodies, suggesting that this compound affects cellular vesicle trafficking systems related to PIN trafficking. The eight chemicals identified here will be a useful tool for understanding the mechanisms of IAA transport in plants. [ABSTRACT FROM PUBLISHER]

Among the most fascinated properties of the plant hormone auxin is its ability to promote formation of its own directional transport routes. These gradually narrowing auxin channels form from the auxin source toward the sink and involve coordinated, collective polarization of individual cells. Once established, the channels provide positional information, along which new vascular strands form, for example, during organogenesis, regeneration, or leave venation. The main prerequisite of this still mysterious auxin canalization mechanism is a feedback between auxin signaling and its directional transport. This is manifested by auxin-induced re-arrangements of polar, subcellular localization of PIN-FORMED (PIN) auxin exporters. Immanent open questions relate to how position of auxin source and sink as well as tissue context are sensed and translated into tissue polarization and how cells communicate to polarize coordinately. Recently, identification of the first molecular players opens new avenues into molecular studies of this intriguing example of self-organizing plant development. [ABSTRACT FROM AUTHOR]