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Auxin plays a pivotal role in plant development by activating AUXIN RESPONSE FACTORs (ARFs). Under low auxin levels, ARF activity is inhibited by interacting with Aux/IAAs. Aux/IAAs are degraded when the cellular auxin concentration increases, causing the release of ARF inhibition. Here, we show that levels of the ARF5/MONOPTEROS (MP) protein are regulated in a cell-type-specific and isoform-dependent manner. We find that the stability of MP isoforms is differentially controlled depending on the auxin level. The canonical MP isoform is degraded by the proteasome in root tissues with low auxin levels. While auxin sharpens the MP localization domain in roots, it does not do so in ovules or embryos. Our research highlights a mechanism for providing spatial control of auxin signaling capacity. Together with recent advances in understanding the tissue-specific expression and post-transcriptional modification of auxin signaling components, these results provide insights into understanding how auxin can elicit so many distinct responses., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2024 The Authors. Published by Elsevier Inc. All rights reserved.)

Barley (Hordeum vulgare) is an important global cereal crop and a model in genetic studies. Despite advances in characterising barley genomic resources, few mutant studies have identified genes controlling root architecture and anatomy, which plays a critical role in capturing soil resources. Our phenotypic screening of a TILLING mutant collection identified line TM5992 exhibiting a short-root phenotype compared with wild-type (WT) Morex background. Outcrossing TM5992 with barley variety Proctor and subsequent SNP array-based bulk segregant analysis, fine mapped the mutation to a cM scale. Exome sequencing pinpointed a mutation in the candidate gene HvPIN1a, further confirming this by analysing independent mutant alleles. Detailed analysis of root growth and anatomy in Hvpin1a mutant alleles exhibited a slower growth rate, shorter apical meristem and striking vascular patterning defects compared to WT. Expression and mutant analyses of PIN1 members in the closely related cereal brachypodium (Brachypodium distachyon) revealed that BdPIN1a and BdPIN1b were redundantly expressed in root vascular tissues but only Bdpin1a mutant allele displayed root vascular defects similar to Hvpin1a. We conclude that barley PIN1 genes have sub-functionalised in cereals, compared to Arabidopsis (Arabidopsis thaliana), where PIN1a sequences control root vascular patterning., (© 2024 The Author(s). New Phytologist © 2024 New Phytologist Foundation.)

Studying the developmental genetics of plant organs requires following gene expression in specific tissues. To facilitate this, we have developed dual expression anatomy lines, which incorporate a red plasma membrane marker alongside a fluorescent reporter for a gene of interest in the same vector. Here, we adapted the GreenGate cloning vectors to create two destination vectors showing strong marking of cell membranes in either the whole root or specifically in the lateral roots. This system can also be used in both embryos and whole seedlings. As proof of concept, we follow both gene expression and anatomy in Arabidopsis (Arabidopsis thaliana) during lateral root organogenesis for a period of over 24 h. Coupled with the development of a flow cell and perfusion system, we follow changes in activity of the DII auxin sensor following application of auxin., (© The Author(s) 2021. Published by Oxford University Press on behalf of American Society of Plant Biologists.)

During plant development, a precise balance of cytokinin is crucial for correct growth and patterning, but it remains unclear how this is achieved across different cell types and in the context of a growing organ. Here we show that in the root apical meristem, the TMO5/LHW complex increases active cytokinin levels via two cooperatively acting enzymes. By profiling the transcriptomic changes of increased cytokinin at single-cell level, we further show that this effect is counteracted by a tissue-specific increase in CYTOKININ OXIDASE 3 expression via direct activation of the mobile transcription factor SHORTROOT. In summary, we show that within the root meristem, xylem cells act as a local organizer of vascular development by non-autonomously regulating cytokinin levels in neighbouring procambium cells via sequential induction and repression modules., (© 2021. The Author(s), under exclusive licence to Springer Nature Limited.)

Root branching is influenced by the soil environment and exhibits a high level of plasticity. We report that the radial positioning of emerging lateral roots is influenced by their hydrological environment during early developmental stages. New lateral root primordia have both a high degree of flexibility in terms of initiation and development angle towards the available water. Our observations reveal how the external hydrological environment regulates lateral root morphogenesis.

Pattern formation is typically controlled through the interaction between molecular signals within a given tissue. During early embryonic development, roots of the model plant Arabidopsis thaliana have a radially symmetric pattern, but a heterogeneous input of the hormone auxin from the two cotyledons forces the vascular cylinder to develop a diarch pattern with two xylem poles. Molecular analyses and mathematical approaches have uncovered the regulatory circuit that propagates this initial auxin signal into a stable cellular pattern. The diarch pattern seen in Arabidopsis is relatively uncommon among flowering plants, with most species having between three and eight xylem poles. Here, we have used multiscale mathematical modelling to demonstrate that this regulatory module does not require a heterogeneous auxin input to specify the vascular pattern. Instead, the pattern can emerge dynamically, with its final form dependent upon spatial constraints and growth. The predictions of our simulations compare to experimental observations of xylem pole number across a range of species, as well as in transgenic systems in Arabidopsis in which we manipulate the size of the vascular cylinder. By considering the spatial constraints, our model is able to explain much of the diversity seen in different flowering plant species., Competing Interests: Competing interestsThe authors declare no competing or financial interests., (© 2019. Published by The Company of Biologists Ltd.)

Plants adapt to heterogeneous soil conditions by altering their root architecture. For example, roots branch when in contact with water by using the hydropatterning response. We report that hydropatterning is dependent on auxin response factor ARF7. This transcription factor induces asymmetric expression of its target gene LBD16 in lateral root founder cells. This differential expression pattern is regulated by posttranslational modification of ARF7 with the small ubiquitin-like modifier (SUMO) protein. SUMOylation negatively regulates ARF7 DNA binding activity. ARF7 SUMOylation is required to recruit the Aux/IAA (indole-3-acetic acid) repressor protein IAA3. Blocking ARF7 SUMOylation disrupts IAA3 recruitment and hydropatterning. We conclude that SUMO-dependent regulation of auxin response controls root branching pattern in response to water availability., (Copyright © 2018 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works.)

Phosphate (P) is an essential macronutrient for plant growth. Roots employ adaptive mechanisms to forage for P in soil. Root hair elongation is particularly important since P is immobile. Here we report that auxin plays a critical role promoting root hair growth in Arabidopsis in response to low external P. Mutants disrupting auxin synthesis (taa1) and transport (aux1) attenuate the low P root hair response. Conversely, targeting AUX1 expression in lateral root cap and epidermal cells rescues this low P response in aux1. Hence auxin transport from the root apex to differentiation zone promotes auxin-dependent hair response to low P. Low external P results in induction of root hair expressed auxin-inducible transcription factors ARF19, RSL2, and RSL4. Mutants lacking these genes disrupt the low P root hair response. We conclude auxin synthesis, transport and response pathway components play critical roles regulating this low P root adaptive response.

The Arabidopsis root has provided an excellent model for understanding patterning processes and cell fate specification. Vascular patterning represents an especially interesting process, as new positional information must be generated to transform an approximately radially symmetric root pole into a bisymmetric structure with a single xylem axis. This process requires both growth of the embryonic tissue alongside the subsequent patterning. Recently researchers have identified a series of transcription factors that modulate cell divisions to control vascular tissues growth. Spatial regulation in the signalling of two hormones, auxin and cytokinin, combine with other transcription factors to pattern the xylem axis. We are now witnessing the discovery of increasingly complex interactions between these hormones that can be interpreted through the use of mathematical models., (Copyright © 2017 The Authors. Published by Elsevier Ltd.. All rights reserved.)

The root vascular tissues provide an excellent system for studying organ patterning, as the specification of these tissues signals a transition from radial symmetry to bisymmetric patterns. The patterning process is controlled by the combined action of hormonal signaling/transport pathways, transcription factors, and miRNA that operate through a series of non-linear pathways to drive pattern formation collectively. With the discovery of multiple components and feedback loops controlling patterning, it has become increasingly difficult to understand how these interactions act in unison to determine pattern formation in multicellular tissues. Three independent mathematical models of root vascular patterning have been formulated in the last few years, providing an excellent example of how theoretical approaches can complement experimental studies to provide new insights into complex systems. In many aspects these models support each other; however, each study also provides its own novel findings and unique viewpoints. Here we reconcile these models by identifying the commonalities and exploring the differences between them by testing how transferable findings are between models. New simulations herein support the hypothesis that an asymmetry in auxin input can direct the formation of vascular pattern. We show that the xylem axis can act as a sole source of cytokinin and specify the correct pattern, but also that broader patterns of cytokinin production are also able to pattern the root. By comparing the three modeling approaches, we gain further insight into vascular patterning and identify several key areas for experimental investigation., (© The Author 2016. Published by Oxford University Press on behalf of the Society for Experimental Biology. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Lateral root primordia (LRP) originate from pericycle stem cells located deep within parental root tissues. LRP emerge through overlying root tissues by inducing auxin-dependent cell separation and hydraulic changes in adjacent cells. The auxin-inducible auxin influx carrier LAX3 plays a key role concentrating this signal in cells overlying LRP. Delimiting LAX3 expression to two adjacent cell files overlying new LRP is crucial to ensure that auxin-regulated cell separation occurs solely along their shared walls. Multiscale modeling has predicted that this highly focused pattern of expression requires auxin to sequentially induce auxin efflux and influx carriers PIN3 and LAX3, respectively. Consistent with model predictions, we report that auxin-inducible LAX3 expression is regulated indirectly by AUXIN RESPONSE FACTOR 7 (ARF7). Yeast one-hybrid screens revealed that the LAX3 promoter is bound by the transcription factor LBD29, which is a direct target for regulation by ARF7. Disrupting auxin-inducible LBD29 expression or expressing an LBD29-SRDX transcriptional repressor phenocopied the lax3 mutant, resulting in delayed lateral root emergence. We conclude that sequential LBD29 and LAX3 induction by auxin is required to coordinate cell separation and organ emergence., (© 2016. Published by The Company of Biologists Ltd.)

Asymmetric distribution of the hormone auxin organizes plant cell fate and drives specification of new organs. Such asymmetries are regulated by polarized auxin transporters called PINs. But what controls PIN polarity? A new study shows that another hormone, cytokinin, degrades PINs on specific membranes to direct auxin flux., (Copyright © 2014 Elsevier Ltd. All rights reserved.)

As multicellular organisms grow, positional information is continually needed to regulate the pattern in which cells are arranged. In the Arabidopsis root, most cell types are organized in a radially symmetric pattern; however, a symmetry-breaking event generates bisymmetric auxin and cytokinin signaling domains in the stele. Bidirectional cross-talk between the stele and the surrounding tissues involving a mobile transcription factor, SHORT ROOT (SHR), and mobile microRNA species also determines vascular pattern, but it is currently unclear how these signals integrate. We use a multicellular model to determine a minimal set of components necessary for maintaining a stable vascular pattern. Simulations perturbing the signaling network show that, in addition to the mutually inhibitory interaction between auxin and cytokinin, signaling through SHR, microRNA165/6, and PHABULOSA is required to maintain a stable bisymmetric pattern. We have verified this prediction by observing loss of bisymmetry in shr mutants. The model reveals the importance of several features of the network, namely the mutual degradation of microRNA165/6 and PHABULOSA and the existence of an additional negative regulator of cytokinin signaling. These components form a plausible mechanism capable of patterning vascular tissues in the absence of positional inputs provided by the transport of hormones from the shoot.

In Arabidopsis, lateral roots originate from pericycle cells deep within the primary root. New lateral root primordia (LRP) have to emerge through several overlaying tissues. Here, we report that auxin produced in new LRP is transported towards the outer tissues where it triggers cell separation by inducing both the auxin influx carrier LAX3 and cell-wall enzymes. LAX3 is expressed in just two cell files overlaying new LRP. To understand how this striking pattern of LAX3 expression is regulated, we developed a mathematical model that captures the network regulating its expression and auxin transport within realistic three-dimensional cell and tissue geometries. Our model revealed that, for the LAX3 spatial expression to be robust to natural variations in root tissue geometry, an efflux carrier is required--later identified to be PIN3. To prevent LAX3 from being transiently expressed in multiple cell files, PIN3 and LAX3 must be induced consecutively, which we later demonstrated to be the case. Our study exemplifies how mathematical models can be used to direct experiments to elucidate complex developmental processes.

In Arabidopsis thaliana, lateral roots (LRs) initiate from anticlinal cell divisions of pericycle founder cells. The formation of LR primordia is regulated antagonistically by the phytohormones cytokinin and auxin. It has previously been shown that cytokinin has an inhibitory effect on the patterning events occurring during LR formation. However, the molecular players involved in cytokinin repression are still unknown. In a similar manner to protoxylem formation in Arabidopsis roots, in which AHP6 (ARABIDOPSIS HISTIDINE PHOSPHOTRANSFER PROTEIN 6) acts as a cytokinin inhibitor, we reveal that AHP6 also functions as a cytokinin repressor during early stages of LR development. We show that AHP6 is expressed at different developmental stages during LR formation and is required for the correct orientation of cell divisions at the onset of LR development. Moreover, we demonstrate that AHP6 influences the localization of the auxin efflux carrier PIN1, which is necessary for patterning the LR primordia. In summary, we show that the inhibition of cytokinin signaling through AHP6 is required to establish the correct pattern during LR initiation.

The plant hormone cytokinin controls root growth by balancing the division and differentiation of stem cells. But what controls accumulation of cytokinin? A new study has identified a regulatory loop between a transcription factor, PHABULOSA, and cytokinin biosynthesis that creates robust domains of cytokinin activity., (Copyright © 2012 Elsevier Ltd. All rights reserved.)

In the shoot pole of Arabidopsis embryos, radial symmetry is broken by cotyledon specification. Subsequently, the radial pattern of the embryo axis is converted to bisymmetric. In a recent publication, we showed that distinct boundaries of hormonal signalling output specify the vascular pattern in the root meristem through a mutually inhibitory feedback loop between the hormones auxin and cytokinin. We observed that during embryogenesis, symmetry breakage in the root pole coincided with an influx of auxin from the cotyledons. In this manuscript, we provide genetic data to support the role of the cotyledons in initiating symmetry breaking in the embryonic root pole. Mutants with alterations in cotyledon number fail to establish bisymmetry in the embryo axis. These data further support the idea that input from the cotyledons may be required for the propagation of bisymmetry from the cotyledons to the embryonic root.

Despite their relatively simple appearance, roots are incredibly complex organs that are highly adapted to differing environments. Many aspects of root development are co-ordinated by subtle spatial differences in the concentrations of the phytohormones auxin and cytokinin. Events from the formation of a root during embryogenesis to the determination of the network of lateral roots are controlled by interactions between these hormones. Recently, interactions have been defined where auxin signaling promotes the expression of cytokinin signaling inhibitors, cytokinin signaling promotes the expression of auxin signaling inhibitors and finally where cytokinin signaling regulates the complex network of auxin transport proteins to position zones of high auxin signaling. We are witnessing a period of discovery in which we are beginning to understand how these hormonal pathways communicate to regulate root formation., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

How are the lateral root founder cells specified in the pericycle to initiate lateral root development? An Aux/IAA28 signaling module activates transcription factor GATA23 to control founder cell identity., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

The cytokinin class of phytohormones regulates division and differentiation of plant cells. They are perceived and signaled by a phosphorelay mechanism similar to those observed in prokaryotes. Research into the components of phosphorelay had previously been marred by genetic redundancy. However, recent studies have addressed this with the creation of high-order mutants. In addition, several new elements regulating cytokinin signaling have been identified. This has uncovered many roles in diverse developmental and physiological processes. In this review, we look at these processes specifically in the context of root development. We focus on the formation and maintenance of the root apical meristem, primary and secondary vascular development, lateral root emergence and development, and root nodulation. We believe that the root is an ideal organ with which to investigate cytokinin signaling in a wider context.

The cell lineages that form the transporting tissues (xylem and phloem) and the intervening pluripotent procambial tissue originate from stem cells near the root tip. We demonstrate that in Arabidopsis, cytokinin phytohormones negatively regulate protoxylem specification. AHP6, an inhibitory pseudophosphotransfer protein, counteracts cytokinin signaling, allowing protoxylem formation. Conversely, cytokinin signaling negatively regulates the spatial domain of AHP6 expression. Thus, by controlling the identity of cell lineages, the reciprocal interaction of cytokinin signaling and its spatially specific modulator regulates proliferation and differentiation of cell lineages during vascular development, demonstrating a previously unrecognized regulatory circuit underlying meristem organization.

Phosphorus is a crucial macronutrient for plants playing a critical role in many cellular signaling and energy cycling processes. In light of this, phosphorus acquisition efficiency is an important target trait for crop improvement, but it also provides an ecological adaptation for growth of plants in low nutrient environments. Increased root hair density has been shown to improve phosphorus uptake and plant health in a number of species. In several plant families, including Brassicaceae, root hair bearing cells are positioned on the epidermis according to their position in relation to cortex cells, with hair cells positioned in the cleft between two underlying cortex cells. Thus the number of cortex cells determines the number of epidermal cells in the root hair position. Previous research has associated phosphorus-limiting conditions with an increase in the number of cortex cell files in Arabidopsis thaliana roots, but they have not investigated the spatial or temporal domains in which these extra divisions occur or explored the consequences this has had on root hair formation. In this study, we use 3D reconstructions of rootmeristems to demonstrate that the radial anticlinal cell divisions seen under low phosphate are exclusive to the cortex. When grown on media containing replete levels of phosphorous, A. thaliana plants almost invariably show eight cortex cells; however when grown in phosphate limited conditions, seedlings develop up to 16 cortex cells (with 10-14 being the most typical). This results in a significant increase in the number of epidermal cells at hair forming positions. These radial anticlinal divisions occur within the initial cells and can be seen within 24 h of transfer of plants to low phosphorous conditions. We show that these changes in the underlying cortical cells feed into epidermal patterning by altering the regular spacing of root hairs. [ABSTRACT FROM AUTHOR]

The article discusses a study by M. Taylor-Teeples and colleagues on the role of genetic regulatory network in controlling the biopolymer synthesis in root xylem cells of Arabidopsis thaliana. Topics cited include the screening of transcriptor factor expression in roots to determine their binding ability in genes encoding cell wall components, effect of lignin on the extraction of sugars in cellulose and hemicelluloses, and description of feedforward loops formation in transcription factors.

Summary The cultivation of many crops relies on the formation of chimeric plants, where roots from one variety are grafted onto the shoot of another. A new study uncovers how two plants connect and demonstrates that the root and shoot do not contribute equally to the union. [ABSTRACT FROM AUTHOR]