Root Cell Division Research Articles

Non-thermal plasma enhances growth and salinity tolerance of bok choy (Brassica rapa subsp. chinensis) in hydroponic culture.

In this study, we aimed to examine the growth, physiological and biochemical status, and responses to salinity stress of bok choy (Brassica rapa subsp. chinensis) cultivated in a hydroponic system with a plasma-treated solution. Plasma gas generated using a cylindrical dielectric barrier discharge or air (control) was injected into Hoagland nutrient solution once a week for different durations (0, 5, and 10 min). After 4 weeks, the length of the shoots and roots, number of leaves, and dry weight of bok choy plants significantly increased in individuals grown with Hoagland solution treated with plasma gas for 10 min. An increase in dry weight of individual plants of approximately 80.5% was observed in plants in the plasma-treated group compared to those in a control group. The levels of chlorophyll, total soluble proteins, and nitrogen uptake, and transcription of genes related to salinity stress tolerance-WRKY2, HHP3, and ABI1- were also significantly elevated in bok choy grown with plasma treated Hoagland solution. Moreover, when exposed to 20 mM NaCl, plant length and leaf number were significantly increased, in the group grown with Hoagland solution treated with plasma gas for 10 min. Level of H2O2 was significantly elevated in the treated nutrient solutions. In plants grown with the treated nutrient solution, intracellular NO was highly detected in the cell division and elongation zone of roots. Our findings suggest that plasma treatment of nutrient solutions in hydroponic culture systems may improve the growth, physiological and biochemical status, and tolerance to salinity stress in plants, and a crucial role of H2O2 generated in the treated nutrient solutions may play in this improvement.

Enhancing Solanum lycopersicum Resilience: Bacterial Cellulose Alleviates Low Irrigation Stress and Boosts Nutrient Uptake.

The use of natural-origin biomaterials in bioengineering has led to innovative approaches in agroforestry. Bacterial cellulose (BC), sharing the same chemical formula as plant-origin cellulose (PC), exhibits significantly different biochemical properties, including a high degree of crystallinity and superior water retention capacity. Previous research showed that natural-origin glucose-based chitin enhanced plant growth in both herbaceous and non-herbaceous plants. In this study, we produced BC in the laboratory and investigated its effects on the substrate and on Solanum lycopersicum seedlings. Soil amended with BC increased root growth compared with untreated seedlings. Additionally, under limited irrigation conditions, BC increased global developmental parameters including fresh and dry weight, as well as total carbon and nitrogen content. Under non-irrigation conditions, BC contributed substantially to plant survival. RNA sequencing (Illumina®) on BC-treated seedlings revealed that BC, despite its bacterial origin, did not stress the plants, confirming its innocuous nature, and it lightly induced genes related to root development and cell division as well as inhibition of stress responses and defense. The presence of BC in the organic substrate increased soil availability of phosphorus (P), iron (Fe), and potassium (K), correlating with enhanced nutrient uptake in plants. Our results demonstrate the potential of BC for improving soil nutrient availability and plant tolerance to low irrigation, making it valuable for agricultural and forestry purposes in the context of global warming.

Circadian Rhythm and Nitrogen Metabolism Participate in the Response of Boron Deficiency in the Root of Brassica napus.

Boron (B) deficiency has been shown to inhibit root cell growth and division. However, the precise mechanism underlying B deficiency-mediated root tip growth inhibition remains unclear. In this study, we investigated the role of BnaA3.NIP5;1, a gene encoding a boric acid channel, in Brassica napus (B. napus). BnaA3.NIP5;1 is expressed in the lateral root cap and contributes to B acquisition in the root tip. Downregulation of BnaA3.NIP5;1 enhances B sensitivity in B. napus, resulting in reduced shoot biomass and impaired root tip development. Transcriptome analysis was conducted on root tips from wild-type B. napus (QY10) and BnaA3.NIP5;1 RNAi lines to assess the significance of B dynamics in meristematic cells during seedling growth. Differentially expressed genes (DEGs) were significantly enriched in plant circadian rhythm and nitrogen (N) metabolism pathways. Notably, the circadian-rhythm-related gene HY5 exhibited a similar B regulation pattern in Arabidopsis to that observed in B. napus. Furthermore, Arabidopsis mutants with disrupted circadian rhythm (hy5/cor27/toc1) displayed heightened sensitivity to low B compared to the wild type (Col-0). Consistent with expectations, B deficiency significantly disrupted N metabolism in B. napus roots, affecting nitrogen concentration, nitrate reductase enzyme activity, and glutamine synthesis. Interestingly, this disruption was exacerbated in BnaA3NIP5;1 RNAi lines. Overall, our findings highlight the critical role of B dynamics in root tip cells, impacting circadian rhythm and N metabolism, ultimately leading to retarded growth. This study provides novel insights into B regulation in root tip development and overall root growth in B. napus.

Effect of sewage wastewater on the mitotic index of the root tips of Allium cepa in koya city /Kurdistan region of Iraq

The purpose of the experiment was to investigate onion (Allium cepa) root growth, cell division, and mitotic index. The onions were placed into four different water samples (Control water, Azadi bridge, Sar Razan, and Sar Baskan) for 72 hours in beakers. The results showed that the number of divided cells decreased significantly in the sewage water compared with the control water. The control water had a high percentage of divided cells (18%), whereas the Sar Razan sewage water had the least percentage of divided cells (1.3%). The percentage of divided cells in Sar Baskan sewage water was 2%, and the percentage of divided cells in Azadi Bridge was 2.6%. It was found that the mitotic index and the number of divided cells increased in the control water compared to the sewage water. The results showed that sewage water has a greater impact on root growth, cell division, and mitotic index of root tips of Allium cepa.

Ethylene and ROS mediate root growth inhibition induced by the endocrine disruptor bisphenol A (BPA)

Bisphenol A (BPA) functions as a detrimental substance that disrupts the endocrine system in animals while also impeding the growth and development of plants. In our previous study, we demonstrated that BPA hinders the growth of roots in Arabidopsis by diminishing cell division and elongation, which is ascribed to the increased accumulation and redistribution of auxin. Here, we examined the mediation of ROS and ethylene in BPA-induced auxin accumulation and root growth inhibition. BPA enhanced ROS levels, and ROS increased auxin contents but reduced cell division activity and the expression of EXPA8 involved in root elongation. ROS scavenger treatment reversed BPA-triggered root growth retardation, auxin accumulation, and cell division inhibition. In addition, BPA induced ethylene, and ethylene synthesis inhibitor treatment reversed BPA-triggered root growth retardation and auxin accumulation. Taken together, ROS and ethylene are involved in BPA-inhibited cell elongation and cell division by mediating auxin accumulation and redistribution.

The response of Arabidopsis to the apocarotenoid β-cyclocitric acid reveals a role for SIAMESE-RELATED 5 in root development and drought tolerance

Abstract New regulatory functions in plant development and environmental stress responses have recently emerged for a number of apocarotenoids produced by enzymatic or nonenzymatic oxidation of carotenoids. β-Cyclocitric acid (β-CCA) is one such compound derived from β-carotene, which triggers defense mechanisms leading to a marked enhancement of plant tolerance to drought stress. We show here that this response is associated with an inhibition of root growth affecting both root cell elongation and division. Remarkably, β-CCA selectively induced cell cycle inhibitors of the SIAMESE-RELATED (SMR) family, especially SMR5, in root tip cells. Overexpression of the SMR5 gene in Arabidopsis induced molecular and physiological changes that mimicked in large part the effects of β-CCA. In particular, the SMR5 overexpressors exhibited an inhibition of root development and a marked increase in drought tolerance which is not related to stomatal closure. SMR5 up-regulation induced changes in gene expression that strongly overlapped with the β-CCA–induced transcriptomic changes. Both β-CCA and SMR5 led to a down-regulation of many cell cycle activators (cyclins, cyclin-dependent kinases) and a concomitant up-regulation of genes related to water deprivation, cellular detoxification, and biosynthesis of lipid biopolymers such as suberin and lignin. This was correlated with an accumulation of suberin lipid polyesters in the roots and a decrease in nonstomatal leaf transpiration. Taken together, our results identify the β-CCA–inducible and drought-inducible SMR5 gene as a key component of a stress-signaling pathway that reorients root metabolism from growth to multiple defense mechanisms leading to drought tolerance.

Immunolocalization of Jasmonates and Auxins in Pea Roots in Connection with Inhibition of Root Growth under Salinity Conditions.

Inhibition of root elongation is an important growth response to salinity, which is thought to be regulated by the accumulation of jasmonates and auxins in roots. Nevertheless, the mechanisms of the interaction of these hormones in the regulation of the growth response to salinity are still not clear enough. Their better understanding depends on the study of the distribution of jasmonates and auxins between root cells. This was achieved with the help of immunolocalization of auxin (indoleacetic acid) and jasmonates on the root sections of pea plants. Salinity inhibited root elongation and decreased the size of the meristem zone and the length of cells in the elongation zone. Immunofluorescence based on the use of appropriate, specific antibodies that recognize auxins and jasmonates revealed an increased abundance of both hormones in the meristem zone. The obtained data suggests the participation of either auxins or jasmonates in the inhibition of cell division, which leads to a decrease in the size of the meristem zone. The level of only auxin and not jasmonate increased in the elongation zone. However, since some literature evidence argues against inhibition of root cell division by auxins, while jasmonates have been shown to inhibit this process, we came to the conclusion that elevated jasmonate is a more likely candidate for inhibiting root meristem activity under salinity conditions. Data suggests that auxins, not jasmonates, reduce cell size in the elongation zone of salt-stressed plants, a suggestion supported by the known ability of auxins to inhibit root cell elongation.

Enhanced meristem development, tolerance to oxidative stressand hyposensitivity to nitric oxide in the hypermorphic vq10-H mutant in AtVQ10 gene.

Nitric oxide (NO) and reactive oxygen are common factors in multiple plant responses to stress, and their involvement in hypoxia-triggered responses is key to ensure growth under adverse environmental conditions. Here, we analyse the regulatory functions exerted by hypoxia-, NO-and oxidative stress-inducible Arabidopsis gene coding for the VQ motif-containing protein 10 (VQ10). A hypermorphic vq10-H mutant allowed identifying VQ10-exerted regulation on root and shoot development as well as its role in regulating responses to NO and oxidative stress. Enhanced VQ10 expression in vq10-H plants led to enhanced elongation of the primary root, and increased root cell division and meristem size during early postgermination development. In shoots, VQ10 activation of cell division was counteracted by WRKY33-exerted repression, thus leading to a dwarf bushy phenotype in plants with enhanced VQ10 expression in a wrky33 knock-out background. Low number of differentially expressed genes were identified when vq10-H versus Col-0 plants were compared either under normoxia or hypoxia. vq10-H and VQ10ox plants displayed less tolerance to submergence but, in turn, were more tolerant to oxidative stress and less sensitive to NO than wild-type plants. VQ10 could be a node integrating redox-related regulation on development and stress responses.

Combination of Iron and Zinc Enhanced the Root Cell Division, Mitotic Regularity and Nucleolar Activity of Hexaploid Triticale.

Hexaploid triticale results from crosses between durum wheat and rye. Despite its high agronomic potential, triticale is mainly used for livestock feed. Triticale surpasses their parental species in adaptability and tolerance to abiotic and biotic stresses, being able to grow in acidic soils where a high amount of iron (Fe) and zinc (Zn) is typical. On the other hand, high amounts of these essential trace elements can be cytotoxic to bread wheat. The cytotoxicity induced by seed priming with a high concentration of Fe and Zn impaired root cell division and induced nucleolar changes in bread wheat. Such cytogenetic approaches were expedited and successfully determined cytotoxic and suited micronutrient dosages for wheat nutripriming. With this study, we intended to analyse the hexaploid triticale cv 'Douro' root mitotic cell cycle and nucleolar activity after seed priming performed with aqueous solutions of iron (Fe) and/or zinc (Zn), containing a concentration that was previously considered cytotoxic, to bread wheat and to infer the higher tolerance of triticale to these treatments. The overall cytogenetic data allowed us to conclude that the Fe + Zn treatment enhanced the root mitotic index (MI), mitosis regularity and nucleolar activity of 'Douro' relative to the control and the individual treatments performed with Fe or Zn alone. The Fe + Zn treatment might suit triticale biofortification through seed priming.

Auxin-cytokinin interplay shapes root functionality under low-temperature stress.

Low-temperature stress alters root system architecture. In particular, changes in the levels and response to auxin and cytokinin determine the fate of root architecture and function under stress because of their vital roles in regulating root cell division, differentiation, and elongation. An intricate nexus of genes encoding components of auxin and cytokinin biosynthesis, signaling, and transport components operate to counteract stress and facilitate optimum development. We review the role of auxin transport and signaling and its regulation by cytokinin during root development and stem cell maintenance under low-temperature stress. We highlight intricate mechanisms operating in root stem cells to minimize DNA damage by altering phytohormone levels, and discuss a working model for cytokinin in low-temperatures stress response.

Autophagy-Mediated Phosphate Homeostasis in Arabidopsis Involves Modulation of Phosphate Transporters

Autophagy in plants is regulated by diverse signaling cascades in response to environmental changes. Fine-tuning of its activity is critical for the maintenance of cellular homeostasis under basal and stressed conditions. In this study, we compared the Arabidopsis autophagy-related (ATG) system transcriptionally under inorganic phosphate (Pi) deficiency versus nitrogen deficiency and showed that most ATG genes are only moderately upregulated by Pi starvation, with relatively stronger induction of AtATG8f and AtATG8h among the AtATG8 family. We found that Pi shortage increased the formation of GFP-ATG8f-labeled autophagic structures and the autophagic flux in the differential zone of the Arabidopsis root. However, the proteolytic cleavage of GFP-ATG8f and the vacuolar degradation of endogenous ATG8 proteins indicated that Pi limitation does not drastically alter the autophagic flux in the whole roots, implying a cell type-dependent regulation of autophagic activities. At the organismal level, the Arabidopsis atg mutants exhibited decreased shoot Pi concentrations and smaller meristem sizes under Pi sufficiency. Under Pi limitation, these mutants showed enhanced Pi uptake and impaired root cell division and expansion. Despite a reduced steady-state level of several PHOSPHATE TRANSPORTER 1s (PHT1s) in the atg root, cycloheximide treatment analysis suggested that the protein stability of PHT1;1/2/3 is comparable in the Pi-replete wild type and atg5-1. By contrast, the degradation of PHT1;1/2/3 is enhanced in the Pi-deplete atg5-1. Our findings reveal that both basal autophagy and Pi starvation-induced autophagy are required for the maintenance of Pi homeostasis and may modulate the expression of PHT1s through different mechanisms.

Impact of Fire Recurrence and Induced Water Stress on Seed Germination and Root Mitotic Cell Cycle of Pinus pinaster Aiton

Climate change will increase the frequency of drought, heat waves, and wildfires. We intended to analyse how fire recurrence and/or induced water stress can affect seed germination and root cell division in Pinus pinaster Aiton. Seeds from stands with no prior fire history and from post-fire regeneration (in areas burnt once, twice, and thrice) in northern Portugal were germinated in distilled water (control) and polyethylene glycol (PEG) to simulate water stress for four weeks, followed by a recovery period. Roots were analysed cytogenetically. The germination index of the Pinus pinaster seeds was not statistically influenced by the induction of osmotic stress, nor by the fire recurrence of the stands. The mean germination time (MGT) was 10–29 days and 1–36 days for the stress and recovery periods, respectively, and increased with PEG concentration. The 20% PEG treatment inhibited root growth after germination. The 10% PEG treatment induced a high frequency of cytogenetic anomalies, mostly in the sites which experienced fire exposure. While fire recurrence did not affect the germination rate, it seemed to reduce the water stress response, negatively impacting cell division and impair root growth.

A novel type of phytosulfokine, PSK-ε, positively regulates root elongation and formation of lateral roots and root nodules in Medicago truncatula

ABSTRACT Phytosulfokines (PSKs) are a class of tyrosine-sulfated pentapeptides. PSK-α, PSK-γ, and PSK-δ are three reported PSK members involved in regulating plant growth, development, and resistance to biotic and abiotic stresses. Here, we reported a novel type of PSK, PSK-ε with the sequence YSO3VYSO3TN, and its precursor proteins (MtPSKε, LjPSKε, and GmPSKε), specifically from legume species. PSK-ε peptide differs from PSK-δ by one amino acid and is close to PSK-δ in the phylogenetic relationship. Expression profile analysis showed that MtPSKε was highly expressed in Medicago truncatula roots, especially in root tips and emerged lateral roots. Application of the synthetic sulfated PSK-ε peptide and overexpression of MtPSKε significantly promoted M. truncatula root elongation and increased lateral root number, probably by inducing cell division and expansion in roots. Furthermore, MtPSKε expression was induced by rhizobia infection and was detected in root nodules including nodule primordia. Both PSK-ε peptide treatment and MtPSKε overexpression significantly increased nodule number in M. truncatula. Taken together, these results demonstrate that PSK-ε, a novel type of phytosulfokine, positively regulates root elongation and formation of lateral root and root nodule in M. truncatula.

Prospecting toxicity of octocrylene in Allium cepa L. and Eisenia fetida Sav.

Octocrylene sunscreen is found in different environmental compartments. Unlike aquatic organisms, there are few studies evaluating the adverse effects caused by this pollutant on terrestrial plants, and no studies on soil fauna. In this study, octocrylene was evaluated at concentrations of 10, 100, and 1000µg/L for phytotoxicity, cytogenotoxicity, and oxidative stress in Allium cepa L., and acute toxicity and oxidative stress in Eisenia fetida Sav. In A. cepa, at concentrations of 100 and 1000µg/L, octocrylene reduced the germination potential in seeds, inhibited root elongation, and caused disturbance in cell division in roots. In E. fetida, the concentration of 1000µg/L promoted an avoidance rate of 80%, while 10µg/L caused a hormesis effect. The concentrations 100 and 1000µg/L caused lipid peroxidation in A. cepa and E. fetida. Based on the results, the recurrent use of biosolids in soil fertilization, as well as the irrigation of plants with wastewater, with the presence of octocrylene can negatively impact the survival of different species that depend directly or indirectly on the soil.

Comparative Transcriptome Analysis of Deep-Rooting and Shallow-Rooting Potato (Solanum tuberosum L.) Genotypes under Drought Stress.

The selection and breeding of deep rooting and drought-tolerant varieties has become a promising approach for improving the yield and adaptability of potato (Solanum tuberosum L.) in arid and semiarid areas. Therefore, the discovery of root-development-related genes and drought tolerance signaling pathways in potato is important. In this study, we used deep-rooting (C119) and shallow-rooting (C16) potato genotypes, with different levels of drought tolerance, to achieve this objective. Both genotypes were treated with 150 mM mannitol for 0 h (T0), 2 h (T2), 6 h (T6), 12 h (T12), and 24 h (T24), and their root tissues were subjected to comparative transcriptome analysis. A total of 531, 1571, 1247, and 3540 differentially expressed genes (DEGs) in C16 and 1531, 1108, 674, and 4850 DEGs in C119 were identified in T2 vs. T0, T6 vs. T2, T12 vs. T6, and T24 vs. T12 comparisons, respectively. Gene expression analysis indicated that a delay in the onset of drought-induced transcriptional changes in C16 compared with C119. Functional enrichment analysis revealed genotype-specific biological processes involved in drought stress tolerance. The metabolic pathways of plant hormone transduction and MAPK signaling were heavily involved in the resistance of C16 and C119 to drought, while abscisic acid (ABA), ethylene, and salicylic acid signal transduction pathways likely played more important roles in C119 stress responses. Furthermore, genes involved in root cell elongation and division showed differential expression between the two genotypes under drought stress. Overall, this study provides important information for the marker-assisted selection and breeding of drought-tolerant potato genotypes.

MITOGEN-ACTIVATED PROTEIN KINASE PHOSPHATASE 1 mediates root sensing of serotonin through jasmonic acid signaling and modulating reactive oxygen species

Serotonin (5-hydroxytryptamine) acts as a neurotransmitter in mammals and is widely distributed in the plant kingdom, where it influences root growth and defense. Mitogen-Activated Protein Kinases (MAPKs) and MAPK phosphatases (MKPs) play critical functions in decoding hormonal signalling, but their possible roles in mediating serotonin responses await investigation. In this report, we unveiled positive roles for the MITOGEN-ACTIVATED PROTEIN KINASE PHOSPHATASE1 (MKP1) in the inhibition of the primary root growth, cell division, meristem structure, and differentiation events in Arabidopsis seedlings. mkp1 mutants were less sensitive to jasmonic acid applications that halted primary root growth in wild-type (WT) plants, and consistently, the neurotransmitter activated the expression of the JASMONATE ZIM-domain (JAZ) proteins JAZ1 and JAZ10, two critical proteins orchestrating jasmonic acid signalling. This effect correlated with exacerbated production of endogenous reactive oxygen species (ROS) in the WT, a process constitutively manifested in mkp1 mutants. These data help to clarify the relationship between serotonin and growth/defense trade-offs, and reveal the importance of the MAPK pathway in root development through ROS production.

Cyto-Genotoxic Effect Causing Potential of Polystyrene Micro-Plastics in Terrestrial Plants.

The polystyrene micro-plastics (Ps-MPs) is one of the leading pollutants found in both aquatic and terrestrial ecosystems. While most of the studies on the morphology and cyto-toxicity of MPs have been based on aquatic organisms, their effects on terrestrial plants are still scarcely known. The present study was an attempt to measure the effect of different sizes (80, 100, 200, 500, 1000, 2000, 4000, and 8000 nm) and concentrations (100 and 400 mg/L) of Ps-MPs on the root length and chromosomes of root tip cells of Allium cepa using A. cepa root chromosomal aberration assay. Large size Ps-MPs (4000 and 8000 nm) showed the highest reduction in A. cepa root length; however, the differences were not significant (at p ≤ 0.05), with respect to negative control (Milli-Q water). The mitotic index showed both significant size- and concentration-dependent decreases, being the lowest (12.06%) in 100 nm at 100 mg/L concentration, with respect to the control (25.05%). The chromosomal abnormality index (CAI) and nuclear abnormality index (NAI) showed significant decreases, with respect to negative control. In addition, the induction of micro-nucleated cells was also observed in Allium root tip cells, when treated with MPs of all sizes, which can predict direct DNA damage to the plant cells. Hence, we conclude that most of the MP sizes caused cyto-toxic and nuclear damage by adversely impacting the spindle formation and induction of micro-nucleated cells in Allium cepa root tip cells. To the best of our knowledge, this is the first study that showed the effect of considerable size range of Ps-MP sizes on the root length and cell division in plants.

CDC48B facilitates the intercellular trafficking of SHORT-ROOT during radial patterning in roots.

CELL DIVISION CONTROL PROTEIN48 (CDC48) is essential for membrane fusion, protein degradation, and other cellular processes. Here, we revealed the crucial role of CDC48B in regulating periclinal cell division in roots by analyzing the recessive gen1 mutant. We identified the GEN1 gene through map-based cloning and verified that GEN1 encodes CDC48B. gen1 showed severely inhibited root growth, increased periclinal cell division in the endodermis, defective middle cortex (MC) formation, and altered ground tissue patterning in roots. Consistent with these phenotypes, CYCLIND 6;1(CYCD6;1), a periclinal cell division marker, was upregulated in gen1 compared to Col-0. The ratio of SHRpro :SHR-GFP fluorescence in pre-dividing nuclei versus the adjacent stele decreased by 33% in gen1, indicating that the trafficking of SHORT-ROOT (SHR) decreased in gen1 when endodermal cells started to divide. These findings suggest that the loss of function of CDC48B inhibits the intercellular trafficking of SHR from the stele to the endodermis, thereby decreasing SHR accumulation in the endodermis. These findings shed light on the crucial role of CDC48B in regulating periclinal cell division in roots.

Distinct mechanisms orchestrate the contra-polarity of IRK and KOIN, two LRR-receptor-kinases controlling root cell division

In plants, cell polarity plays key roles in coordinating developmental processes. Despite the characterization of several polarly localized plasma membrane proteins, the mechanisms connecting protein dynamics with cellular functions often remain unclear. Here, we introduce a polarized receptor, KOIN, that restricts cell divisions in the Arabidopsis root meristem. In the endodermis, KOIN polarity is opposite to IRK, a receptor that represses endodermal cell divisions. Their contra-polar localization facilitates dissection of polarity mechanisms and the links between polarity and function. We find that IRK and KOIN are recognized, sorted, and secreted through distinct pathways. IRK extracellular domains determine its polarity and partially rescue the mutant phenotype, whereas KOIN’s extracellular domains are insufficient for polar sorting and function. Endodermal expression of an IRK/KOIN chimera generates non-cell-autonomous misregulation of root cell divisions that impacts patterning. Altogether, we reveal two contrasting mechanisms determining these receptors’ polarity and link their polarity to cell divisions in root tissue patterning.

Size and activity of the root meristem: A key for drought resistance and a key model of drought-related signaling.

Plants make many adjustments to their growth and development in response to even small changes in water availability. Under such conditions, root elongation can be actively restricted by stress-related signaling mechanisms. Here we look at how the Arabidopsis thaliana root meristem can be affected by moderate water limitation (low water potential, ψw ). Recent characterization of the clade E Growth-Regulating (EGR) protein phosphatases and Microtubule Associated Stress Protein 1 (MASP1) provides an example of how active restriction of root meristem size allows the plant to downregulate root elongation during low ψw stress. EGR2 protein accumulation in cortex cells of the transition zone at the distal end of the root meristem illustrates how the balance of cell division versus cell expansion signals at this critical location can determine meristem size and root elongation during low ψw . These characteristics of EGRs also raise the question of whether they may also be involved in hydrotropism, and, more broadly, whether hydrotropism is a distinct response or a specific manifestation of more general mechanisms used to adjust root growth under moderate severity low ψw whether or not a gradient of water availability is present. These questions, as well as a better understanding of how specific cell layers (cortex and endodermis) seem to have an outsized role in growth regulation and better understanding the roles of plasma membrane-based signaling and polar-localized proteins in the regulation of root meristem size and cell division activity are key to elucidating the cellular mechanisms that determine root growth behavior during soil drying.