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BackgroundAuxin is a versatile plant hormone with important roles in many essential physiological processes. In recent years, significant progress has been made towards understanding the roles of this hormone in plant growth and development. Recent evidence also points to a less well-known but equally important role for auxin as a mediator of environmental adaptation in plants.ScopeThis review briefly discusses recent findings on how plants utilize auxin signalling and transport to modify their root system architecture when responding to diverse biotic and abiotic rhizosphere signals, including macro- and micro-nutrient starvation, cold and water stress, soil acidity, pathogenic and beneficial microbes, nematodes and neighbouring plants. Stress-responsive transcription factors and microRNAs that modulate auxin- and environment-mediated root development are also briefly highlighted.ConclusionsThe auxin pathway constitutes an essential component of the plant's biotic and abiotic stress tolerance mechanisms. Further understanding of the specific roles that auxin plays in environmental adaptation can ultimately lead to the development of crops better adapted to stressful environments.

Summary Wheat is a staple crop produced in arid and semi‐arid areas worldwide, and its production is frequently compromised by water scarcity. Thus, increased drought tolerance is a priority objective for wheat breeding programmes, and among their targets, the NAC transcription factors have been demonstrated to contribute to plant drought response. However, natural sequence variations in these genes have been largely unexplored for their potential roles in drought tolerance. Here, we conducted a candidate gene association analysis of the stress‐responsive NAC gene subfamily in a wheat panel consisting of 700 varieties collected worldwide. We identified a drought responsive gene, TaSNAC8‐6A, that is tightly associated with drought tolerance in wheat seedlings. Further analysis found that a favourable allele TaSNAC8‐6AIn‐313, carrying an insertion in the ABRE promoter motif, is targeted by TaABFs and confers enhanced drought‐inducible expression of TaSNAC8‐6A in drought‐tolerant genotypes. Transgenic wheat and Arabidopsis TaSNAC8‐6A overexpression lines exhibited enhanced drought tolerance through induction of auxin‐ and drought‐response pathways, confirmed by transcriptomic analysis, that stimulated lateral root development, subsequently improving water‐use efficiency. Taken together, our findings reveal that natural variation in TaSNAC8‐6A and specifically the TaSNAC8‐6AIn‐313 allele strongly contribute to wheat drought tolerance and thus represent a valuable genetic resource for improvement of drought‐tolerant germplasm for wheat production.

Plant development under altered nutritional status and environmental conditions and during attack from invaders is highly regulated by plant hormones at the molecular level by various signaling pathways. Previously, reactive oxygen species (ROS) were believed to be harmful as they cause oxidative damage to cells; however, in the last decade, the essential role of ROS as signaling molecules regulating plant growth has been revealed. Plant roots accumulate relatively high levels of ROS, and thus, maintaining ROS homeostasis, which has been shown to regulate the balance between cell proliferation and differentiation at the root tip, is important for proper root growth. However, when the balance is disturbed, plants are unable to respond to the changes in the surrounding conditions and cannot grow and survive. Moreover, ROS control cell expansion and cell differentiation processes such as root hair formation and lateral root development. In these processes, the transcription factor-mediated gene expression network is important downstream of ROS. Although ROS can independently regulate root growth to some extent, a complex crosstalk occurs between ROS and other signaling molecules. Hormone signals are known to regulate root growth, and ROS are thought to merge with these signals. In fact, the crosstalk between ROS and these hormones has been elucidated, and the central transcription factors that act as a hub between these signals have been identified. In addition, ROS are known to act as important signaling factors in plant immune responses; however, how they also regulate plant growth is not clear. Recent studies have strongly indicated that ROS link these two events. In this review, we describe and discuss the role of ROS signaling in root development, with a particular focus on transcriptional regulation. We also summarize the crosstalk with other signals and discuss the importance of ROS as signaling molecules for plant root development.

In plants, postembryonic formation of new organs helps shape the adult organism. This requires the tight regulation of when and where a new organ is formed and a coordination of the underlying cell divisions. To build a root system, new lateral roots are continuously developing, and this process requires the tight coordination of asymmetric cell division in adjacent pericycle cells. We identified EXPANSIN A1 (EXPA1) as a cell wall modifying enzyme controlling the divisions marking lateral root initiation. Loss of EXPA1 leads to defects in the first asymmetric pericycle cell divisions and the radial swelling of the pericycle during auxin-driven lateral root formation. We conclude that a localized radial expansion of adjacent pericycle cells is required to position the asymmetric cell divisions and generate a core of small daughter cells, which is a prerequisite for lateral root organogenesis.

Main conclusion This study unravels a molecular mechanism of miR156/SPL module in regulating lateral root development and nitrogen uptake in apple. Nitrogen is critical in controlling lateral root development such as the availability or lack of nitrogen nutrients which can affect lateral root formation. The miR156/SPL module plays a pivotal role in regulating many aspects of plant development, including the timing of vegetative phase change, floral induction, shoot branching and root development. However, whether the miR156/SPL module functions in nitrate-mediated apple lateral root development remains largely unknown. Here, we revealed the role of miR156/SPL module in regulating lateral root development and root nitrogen uptake in both apple and Arabidopsis . This finding showed that miR156-targeted transcription factor MdSPL23 not only regulates lateral root development but also root nitrogen uptake. The MdSPL23 is bound to the promoter of MdNLP7 to suppress its expression, thereby negatively regulating nitrogen uptake and inhibiting lateral root development in apple. In a likewise manner, the AtSPL9 involved in lateral root development in Arabidopsis is bound to the MdNLP7 homologous gene AtNLP7 to suppress its expression to regulate nitrate-mediated lateral root development. These results suggest that plants regulate nitrogen-mediated root growth through miR156/SPL modules, and this mechanism might be of universal importance.

• Lateral roots (LRs), which form in the plant postembryonically, determine the architecture of the root system. While negative regulatory factors that inhibit LR formation and are counteracted by auxin exist in the pericycle, these factors have not been characterised. • Here, we report that SHI-RELATED SEQUENCE5 (SRS5) is an intrinsic negative regulator of LR formation and that auxin signalling abolishes this inhibitory effect of SRS5. Whereas LR primordia (LRPs) and LRs were fewer and less dense in SRS5ox and Pro35S:SRS5-GFP plants than in the wild-type, they were more abundant and denser in the srs5-2 loss-of-function mutant. SRS5 inhibited LR formation by directly downregulating the expression of LATERAL ORGAN BOUNDARIES-DOMAIN 16 (LBD16) and LBD29. • Auxin repressed SRS5 expression. Auxin-mediated repression of SRS5 expression was not observed in the arf7-1 arf19-1 double mutant, likely because ARF7 and ARF19 bind to the promoter of SRS5 and inhibit its expression in response to auxin. • Taken together, our data reveal that SRS5 negatively regulates LR formation by repressing the expression of LBD16 and LBD29 and that auxin releases this inhibitory effect through ARF7 and ARF19.

Auxin is essential for the regulation of root system architecture by controlling primary root elongation and lateral root (LR) formation. Exogenous auxin has been reported to inhibit primary root elongation and promote the formation of LRs. In this study, LR formation in the primary root was quantitatively evaluated after exogenous auxin treatment by comparing the effects of auxin on two selected zones elongated either before or after auxin application. We determined two main variables in both zones: the LR density per unit of root length (LRD), and the mean phloem pericycle cell length. The total number of phloem pericycle cells (PPCs) per unit of root length was then calculated. Considering that each LR primordium is initiated from four founder cells (FCs), the percentage of PPCs (%PPC) that behave as FCs in a specific root zone was estimated by dividing the number of pericycle cells by four times the LRD. This index was utilized to describe LR initiation. Root zones elongated in the presence of a synthetic auxin (1-naphthalene acetic acid, NAA) at low concentrations (0.01 μM) showed reduced cell length and increased LRD. However, a high concentration of NAA (0.1 μM) strongly reduced both cell length and LRD. In contrast, both low and high levels of NAA stimulated LRD in zones elongated before auxin application. Analysis of the percentage of FCs in the phloem pericycle in zones elongated in the presence or absence of NAA showed that low concentrations of NAA increased the %PFC, indicating that LR initiation is promoted at new sites; however, high concentrations of NAA elicited a considerable reduction in this variable in zones developed in the presence of auxin. As these zones are composed of short pericycle cells, we propose that short pericycle cells are incapable to participate in LR primordium initiation and that auxin modulated initiation of LRs is linked to pericycle cell length.

In Arabidopsis, lateral root primordia (LRPs) originate from pericycle cells located deep within the parental root and have to emerge through endodermal, cortical, and epidermal tissues. These overlaying tissues place biomechanical constraints on the LRPs that are likely to impact their morphogenesis. This study probes the interplay between the patterns of cell division, organ shape, and overlaying tissues on LRP morphogenesis by exploiting recent advances in live plant cell imaging and image analysis. Our 3D/4D image analysis revealed that early stage LRPs exhibit tangential divisions that create a ring of cells corralling a population of rapidly dividing cells at its center. The patterns of division in the latter population of cells during LRP morphogenesis are not stereotypical. In contrast, statistical analysis demonstrated that the shape of new LRPs is highly conserved. We tested the relative importance of cell division pattern versus overlaying tissues on LRP morphogenesis using mutant and transgenic approaches. The double mutant aurora1 (aur1) aur2 disrupts the pattern of LRP cell divisions and impacts its growth dynamics, yet the new organ's dome shape remains normal. In contrast, manipulating the properties of overlaying tissues disrupted LRP morphogenesis. We conclude that the interaction with overlaying tissues, rather than the precise pattern of divisions, is most important for LRP morphogenesis and optimizes the process of lateral root emergence.

Lateral root development progresses through different steps with, the peptides and receptors involved in each of these steps triggering downstream mechanisms upon peptide perception.

Roots take up phosphorus (P) as inorganic phosphate (Pi). Enhanced root proliferation in Pi-rich patches enables plants to capture the unevenly distributed Pi, but the underlying control of root proliferation remains largely unknown. Here, the role of auxin in this response was investigated in maize (Zea mays). A split-root, hydroponics system was employed to investigate root responses to Pi supply, with one (heterogeneous) or both (homogeneous) sides receiving 0 or 500 μm Pi. Maize roots proliferated in Pi-rich media, particularly with heterogeneous Pi supply. The second-order lateral root number was 3-fold greater in roots of plants receiving a heterogeneous Pi supply than in roots of plants with a homogeneous Pi supply. Root proliferation in a heterogeneous Pi supply was inhibited by the auxin transporter inhibitor 1-N-naphthylphthalamic acid (NPA). The proliferation of lateral roots was accompanied by an enhanced auxin response in the apical meristem and vascular tissues at the root tip, as demonstrated in a DR5::RFP marker line. It is concluded that the response of maize root morphology to a heterogeneous Pi supply is modulated by local signals of Pi availability and systemic signals of plant P nutritional status, and is mediated by auxin redistribution.