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Availability of the essential macronutrient nitrogen in soil plays a critical role in plant growth, development, and impacts agricultural productivity. Plants have evolved different strategies for sensing and responding to heterogeneous nitrogen distribution. Modulation of root system architecture, including primary root growth and branching, is among the most essential plant adaptions to ensure adequate nitrogen acquisition. However, the immediate molecular pathways coordinating the adjustment of root growth in response to distinct nitrogen sources, such as nitrate or ammonium, are poorly understood. Here, we show that growth as manifested by cell division and elongation is synchronized by coordinated auxin flux between two adjacent outer tissue layers of the root. This coordination is achieved by nitrate‐dependent dephosphorylation of the PIN2 auxin efflux carrier at a previously uncharacterized phosphorylation site, leading to subsequent PIN2 lateralization and thereby regulating auxin flow between adjacent tissues. A dynamic computer model based on our experimental data successfully recapitulates experimental observations. Our study provides mechanistic insights broadening our understanding of root growth mechanisms in dynamic environments. SYNOPSIS: Effective soil exploitation by plants depends on rapid adjustment of root systems to available nutrients. Here, root growth adaptation to distinct nitrogen sources is shown to involve modulation of auxin distribution via phosphorylation of the PIN2 auxin carrier. Nitrogen source type (nitrate vs. ammonium) differentially regulates cell elongation and division patterns in Arabidopsis root tip tissues.Nitrogen source type regulates directionality of auxin transport in the root tip.Root growth adjustment to nitrogen source is mediated by PIN2 phosphorylation at S439.PIN2 phosphorylation affects auxin flux via modulation of PIN2 polarized localization at the plasma membrane. [ABSTRACT FROM AUTHOR]

The stem cell niche in the root meristem plays a critical role during the development of plant root systems. The polar cell-to-cell transport generates the distribution maximum of the phytohormone auxin that plays a critical role during growth and patterning of roots. Recently, the transcription factor WOX5 (WUSCHEL-RELATED HOMEOBOX 5) was identified as a master regulator of root stem cell niche patterning. Here, we demonstrate that WOX5 coordinates local auxin production and thus controls spatial restriction and maintenance of the auxin maximum in the quiescent center (QC) of the root. In turn, this root-associated auxin maximum feeds back on WOX5 expression via IAA17-dependent regulation of auxin responses. Both our experimental studies and in silico computer modeling simulations suggest that the feedback circuit between WOX5 activity and auxin signaling maximum is at the heart of auxin-mediated root patterning.In plants, the patterning of stem cell-enriched meristems requires a graded auxin response maximum that emerges from the concerted action of polar auxin transport, auxin biosynthesis, auxin metabolism, and cellular auxin response machinery. However, mechanisms underlying this auxin response maximum-mediated root stem cell maintenance are not fully understood. Here, we present unexpected evidence that WUSCHEL-RELATED HOMEOBOX 5 (WOX5) transcription factor modulates expression of auxin biosynthetic genes in the quiescent center (QC) of the root and thus provides a robust mechanism for the maintenance of auxin response maximum in the root tip. This WOX5 action is balanced through the activity of indole-3-acetic acid 17 (IAA17) auxin response repressor. Our combined genetic, cell biology, and computational modeling studies revealed a previously uncharacterized feedback loop linking WOX5-mediated auxin production to IAA17-dependent repression of auxin responses. This WOX5–IAA17 feedback circuit further assures the maintenance of auxin response maximum in the root tip and thereby contributes to the maintenance of distal stem cell (DSC) populations. Our experimental studies and in silico computer simulations both demonstrate that the WOX5–IAA17 feedback circuit is essential for the maintenance of auxin gradient in the root tip and the auxin-mediated root DSC differentiation. [ABSTRACT FROM PUBLISHER]

Plants develop new organs to adjust their bodies to dynamic changes in the environment. How independent organs achieve anisotropic shapes and polarities is poorly understood. To address this question, we constructed a mechano-biochemical model for Arabidopsis root meristem growth that integrates biologically plausible principles. Computer model simulations demonstrate how differential growth of neighboring tissues results in the initial symmetry-breaking leading to anisotropic root growth. Furthermore, the root growth feeds back on a polar transport network of the growth regulator auxin. Model, predictions are in close agreement with in vivo patterns of anisotropic growth, auxin distribution, and cell polarity, as well as several root phenotypes caused by chemical, mechanical, or genetic perturbations. Our study demonstrates that the combination of tissue mechanics and polar auxin transport organizes anisotropic root growth and cell polarities during organ outgrowth. Therefore, a mobile auxin signal transported through immobile cells drives polarity and growth mechanics to coordinate complex organ development. [ABSTRACT FROM AUTHOR]