Auxin is a key regulator of plant growth and development. Within the root tip, auxin distribution plays a crucial role specifying developmental zones and coordinating tropic responses. Determining how the organ-scale auxin pattern is regulated at the cellular scale is essential to understanding how these processes are controlled. In this study, we developed an auxin transport model based on actual root cell geometries and carrier subcellular localizations. We tested model predictions using the DII-VENUS auxin sensor in conjunction with state-of-the-art segmentation tools. Our study revealed that auxin efflux carriers alone cannot create the pattern of auxin distribution at the root tip and that AUX1/LAX influx carriers are also required. We observed that AUX1 in lateral root cap (LRC) and elongating epidermal cells greatly enhance auxin's shootward flux, with this flux being predominantly through the LRC, entering the epidermal cells only as they enter the elongation zone. We conclude that the nonpolar AUX1/LAX influx carriers control which tissues have high auxin levels, whereas the polar PIN carriers control the direction of auxin transport within these tissues.

2Plants can acclimate by using tropisms to link the direction of growth to 41 environmental conditions. Hydrotropism allows roots to forage for water, a process 42 known to depend on abscisic acid (ABA) but whose molecular and cellular basis 43 remains unclear. Here, we show that hydrotropism still occurs in roots after laser 44 ablation removed the meristem and root cap. Additionally, targeted expression 45 studies reveal that hydrotropism depends on the ABA signalling kinase, SnRK2.2, and 46 the hydrotropism-specific MIZ1, both acting specifically in elongation zone cortical 47 cells. Conversely, hydrotropism, but not gravitropism, is inhibited by preventing 48 differential cell-length increases in the cortex, but not in other cell types. We conclude 49 that root tropic responses to gravity and water are driven by distinct tissue-based 50 mechanisms. In addition, unlike its role in root gravitropism, the elongation zone 51 performs a dual function during a hydrotropic response, both sensing a water 52 potential gradient and subsequently undergoing differential growth. 53 3 Tropic responses are differential growth mechanisms that roots use to explore the 54 surrounding soil efficiently. In general, a tropic response can be divided into several steps, 55 comprising perception, signal transduction, and differential growth. All of these steps have 56 been well characterized for gravitropism, where gravity sensing cells in the columella of the 57 root cap generate a lateral auxin gradient, whilst adjacent lateral root cap cells transport 58 auxin to epidermal cells in the elongation zone, thereby triggering the differential growth that 59 drives bending [1][2][3][4] . In gravi-stimulated roots, the lateral auxin gradient is transported 60 principally by AUX1 and PIN carriers [3][4][5] . 61Compared with gravitropism, the tropic response to asymmetric water availability, i.e., 62 hydrotropism, has been far less studied. Previously, it was reported that surgical removal or 63 ablation of the root cap reduces hydrotropic bending in pea [6][7][8] and Arabidopsis thaliana 9 , 64suggesting that the machinery for sensing moisture gradients resides in the root cap. It has 65 also been reported that hydrotropic bending occurs due to differential growth in the 66 elongation zone 7,10 . However unlike gravitropism, hydrotropism in A. thaliana is independent 67 of AUX1 and PIN-mediated auxin transport 11,12 . Indeed, roots bend hydrotropically in the 68 absence of any redistribution of auxin detectable by auxin-responsive reporters 13,14 . 18,19 . 83However it is unclear whether this broad expression pattern is necessary for MIZ1's function 84 in hydrotropism or whether ABA signal transduction components in general have to be 85 expressed in specific root tip tissues for a hydrotropic response. The present study describes 86 a series of experiments in A. thaliana designed to identify the root tissues essential for a 87 hydrotropic response. We report that MIZ1 and a key ABA signal-transduction component 88SnRK2....

In most organisms, the number and distribution of crossovers that occur during meiosis are tightly controlled. All chromosomes must receive at least one ‘obligatory crossover’ and crossovers are prevented from occurring near one another by ‘crossover interference’. However, the mechanistic basis of this phenomenon of crossover interference has remained mostly mysterious. Using quantitative super-resolution cytogenetics and mathematical modelling, we investigate crossover positioning in the Arabidopsis thaliana wild-type, an over-expressor of the conserved E3 ligase HEI10, and a hei10 heterozygous line. We show that crossover positions can be explained by a predictive, diffusion-mediated coarsening model, in which large, approximately evenly-spaced HEI10 foci grow at the expense of smaller, closely-spaced clusters. We propose this coarsening process explains many aspects of Arabidopsis crossover positioning, including crossover interference. Consistent with this model, we also demonstrate that crossover positioning can be predictably modified in vivo simply by altering HEI10 dosage, with higher and lower dosage leading to weaker and stronger crossover interference, respectively. As HEI10 is a conserved member of the RING finger protein family that functions in the interference-sensitive pathway for crossover formation, we anticipate that similar mechanisms may regulate crossover positioning in diverse eukaryotes.