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In all multicellular organisms, transcriptional networks orchestrate organ development. The Arabidopsis root, with its simple structure and indeterminate growth, is an ideal model for investigating the spatiotemporal transcriptional signatures underlying developmental trajectories. To map gene expression dynamics across root cell types and developmental time, we built a comprehensive, organ-scale atlas at single-cell resolution. In addition to estimating developmental progressions in pseudotime, we employed the mathematical concept of optimal transport to infer developmental trajectories and identify their underlying regulators. To demonstrate the utility of the atlas to interpret new datasets, we profiled mutants for two key transcriptional regulators at single-cell resolution, shortroot and scarecrow. We report transcriptomic and in vivo evidence for tissue trans-differentiation underlying a mixed cell identity phenotype in scarecrow. Our results support the atlas as a rich community resource for unraveling the transcriptional programs that specify and maintain cell identity to regulate spatiotemporal organ development., Competing Interests: Declaration of interests P.N.B. is a member of the Developmental Cell advisory board and is the co-founder and Chair of the Scientific Advisory Board of Hi Fidelity Genetics, a company that works on crop root growth., (Copyright © 2022 The Authors. Published by Elsevier Inc. All rights reserved.)

In plants, salicylic acid (SA) is a hormone that mediates a plant's defense against pathogens. SA also takes an active role in a plant's response to various abiotic stresses, including chilling, drought, salinity, and heavy metals. In addition, in recent years, numerous studies have confirmed the important role of SA in plant morphogenesis. In this review, we summarize data on changes in root morphology following SA treatments under both normal and stress conditions. Finally, we provide evidence for the role of SA in maintaining the balance between stress responses and morphogenesis in plant development, and also for the presence of SA crosstalk with other plant hormones during this process.

The intercellular transport of sugars, nutrients, and small molecules is essential for plant growth, development, and adaptation to environmental changes. Various stresses are known to affect the cell-to-cell molecular trafficking modulated by plasmodesmal permeability. However, the mechanisms of plasmodesmata modification and molecules involved in the phloem unloading process under stress are still not well understood. Here, we show that heat stress reduces the root meristem size and inhibits phloem unloading by inducing callose accumulation at plasmodesmata that connect the sieve element and phloem pole pericycle. Furthermore, we identify the loss-of-function of CALLOSE SYNTHASE 8 ( CalS8 ), which is expressed specifically in the phloem pole pericycle, decreasing the plasmodesmal callose deposition at the interface between the sieve element and phloem pole pericycle and alleviating the suppression at root meristem size by heat stress. Our studies indicate the involvement of callose in the interaction between root meristem growth and heat stress and show that CalS8 negatively regulates the thermotolerance of Arabidopsis roots.

The hormones auxin and cytokinin regulate numerous aspects of plant development and often act as an antagonistic hormone pair. One of the more striking examples of the auxin/cytokinin antagonism involves regulation of the shoot/root growth ratio in which cytokinin promotes shoot and inhibits root growth, whereas auxin does the opposite. Control of the shoot/root growth ratio is essential for the survival of terrestrial plants because it allows growth adaptations to water and mineral nutrient availability in the soil. Because a decrease in shoot growth combined with an increase in root growth leads to survival under drought stress and nutrient limiting conditions, it was not surprising to find that auxin promotes, while cytokinin reduces, drought stress tolerance and nutrient uptake. Recent data show that drought stress and nutrient availability also alter the cytokinin and auxin signaling and biosynthesis pathways and that this stress-induced regulation affects cytokinin and auxin in the opposite manner. These antagonistic effects of cytokinin and auxin suggested that each hormone directly and negatively regulates biosynthesis or signaling of the other. However, a growing body of evidence supports unidirectional regulation, with auxin emerging as the primary regulatory component. This master regulatory role of auxin may not come as a surprise when viewed from an evolutionary perspective.

Plant root growth is indeterminate but continuously responds to environmental changes. We previously reported on the severe root growth defect of a double mutant in bZIP17 and bZIP28 ( bz1728 ) modulating the unfolded protein response (UPR). To elucidate the mechanism by which bz1728 seedlings develop a short root, we obtained a series of bz1728 suppressor mutants, called nobiro , for rescued root growth. We focused here on nobiro6 , which is defective in the general transcription factor component TBP-ASSOCIATED FACTOR 12b (TAF12b). The expression of hundreds of genes, including the bZIP60-UPR regulon, was induced in the bz1728 mutant, but these inductions were markedly attenuated in the bz1728nobiro6 mutant. In view of this, we assigned transcriptional cofactor activity via physical interaction with bZIP60 to NOBIRO6/TAF12b. The single nobiro6 / taf12b mutant also showed an altered sensitivity to endoplasmic reticulum stress for both UPR and root growth responses, demonstrating that NOBIRO6/TAF12b contributes to environment-responsive root growth control through UPR., Competing Interests: The authors declare no competing interest., (Copyright © 2022 the Author(s). Published by PNAS.)

Key Message: This study focused on the role of CLE1-7 peptides as defense mediators, and showed that root-expressed CLE3 functions as a systemic signal to regulate defense-related gene expression in shoots. In the natural environment, plants employ diverse signaling molecules including peptides to defend themselves against various pathogen attacks. In this study, we investigated whether CLAVATA3/EMBRYO SURROUNDING REGION-RELATED (CLE) genes (CLE1-7) respond to biotic stimuli. CLE3 showed significant up-regulation upon treatment with flg22, Pep2, and salicylic acid (SA). Quantitative real-time PCR (qRT-PCR) analysis revealed that CLE3 expression is regulated by the NON-EXPRESSOR OF PR GENES1 (NPR1)-dependent SA signaling and flg22-FLAGELLIN-SENSITIVE 2 (FLS2) signaling pathways. We demonstrated that SA-induced up-regulation of CLE3 in roots was required for activation of WRKY33, a gene involved in the regulation of systemic acquired resistance (SAR), in shoots, suggesting that CLE3 functions as a root-derived signal that regulates the expression of defense-related genes in shoots. Microarray analysis of transgenic Arabidopsis lines overexpressing CLE3 under the control of a β-estradiol-inducible promoter revealed that root-confined CLE3 overexpression affected gene expression in both roots and shoots. Comparison of CLE2- and CLE3-induced genes indicated that CLE2 and CLE3 peptides target a few common but largely distinct downstream genes. These results suggest that root-derived CLE3 is involved in the regulation of systemic rather than local immune responses. Our study also sheds light on the potential role of CLE peptides in long-distance regulation of plant immunity., (© 2022. The Author(s), under exclusive licence to Springer Nature B.V.)

Background: Breadfruit (Artocarpus altilis) is a traditional staple tree crop throughout the tropics. Through interspecific grafting, a dwarf phenotype with over 50% reduction in plant height was identified when marang (Artocarpus odoratissimus) rootstocks were used. However, the molecular mechanism underlying the rootstock-induced breadfruit dwarfing is poorly understood., Results: An RNA-sequencing study of breadfruit scions at 22 months after grafting identified 5409 differentially expressed genes (DEGs) of which 2069 were upregulated and 3339 were downregulated in scion stems on marang rootstocks compared to those on self-graft. The DEGs were predominantly enriched for biological processes involved in carbon metabolism, cell wall organization, plant hormone signal transduction and redox homeostasis. The down-regulation of genes encoding vacuolar acid invertases and alkaline/neutral invertases, was consistent with the decreased activity of both enzymes, accompanying with a higher sucrose but lower glucose and fructose levels in the tissues. Key genes of biosynthetic pathways for amino acids, lipids and cell wall were down regulated, reflecting reduction of sucrose utilisation for stem growth on dwarfing rootstocks. Genes encoding sugar transporters, amino acid transporters, choline transporters, along with large number of potassium channels and aquaporin family members were down-regulated in scion stems on marang rootstocks. Lower activity of plasma membrane H + -ATPase, together with the predominance of genes encoding expansins, wall-associated receptor kinases and key enzymes for biosynthesis and re-modelling of cellulose, xyloglucans and pectins in down-regulated DGEs suggested impairment of cell expansion. Signalling pathways of auxin and gibberellin, along with strigolacton and brassinosteroid biosynthetic genes dominated the down-regulated DEGs. Phenylpropanoid pathway was enriched, with key lignin biosynthetic genes down-regulated, and flavonoid biosynthetic genes upregulated in scions on marang rootstocks. Signalling pathways of salicylic acid, jasmonic acid, ethylene and MAPK cascade were significantly enriched in the upregulated DEGs., Conclusions: Rootstock-induced disruption in pathways regulating nutrient transport, sucrose utilisation, cell wall biosynthesis and networks of hormone transduction are proposed to impair cell expansion and stem elongation, leading to dwarf phenotype in breadfruit scions. The information provides opportunity to develop screening strategy for rootstock breeding and selection for breadfruit dwarfing.

Expression pattern of aluminum (Al) tolerance genes is one of the major determinants of Al avoidance/tolerance within plant cultivars. We have performed transcriptome analysis of two contrasting (Al-tolerant, Disang; Al-sensitive, Joymati) cultivars of India's North Eastern region, an indica rice diversity hotspot, on exposure to excess Al 3+ treatment in acidic condition. Co-expression analysis and SNPs enrichment analysis proposed the role of both trans-acting and cis-acting polymorphisms in Al signaling in the Al-tolerant cultivar. We proposed ten major genes, including arginine decarboxylase, phytase, and beta-glucosidase aggregating factor as candidates responsible for Al tolerance based on transcriptome analysis. Al 3+ stress led to changes in the alternative splicing profile of the Al-tolerant cultivar. These studies demonstrated the transcriptional variations affiliated to Al avoidance/tolerance in contrasting indica rice of North East India and provided us with several candidate genes responsible for Al tolerance.

Rhizobial infection of legume roots during the development of nitrogen-fixing root nodules can occur intracellularly, through plant-derived infection threads traversing cells, or intercellularly, via bacterial entry between epidermal plant cells. Although it is estimated that around 25% of all legume genera are intercellularly infected, the pathways and mechanisms supporting this process have remained virtually unexplored due to a lack of genetically amenable legumes that exhibit this form of infection. In this study, we report that the model legume Lotus japonicus is infected intercellularly by the IRBG74 strain, recently proposed to belong to the Agrobacterium clade of the Rhizobiaceae. We demonstrate that the resources available for L. japonicus enable insight into the genetic requirements and fine-tuning of the pathway governing intercellular infection in this species. Inoculation of L. japonicus mutants shows that Ethylene-responsive factor required for nodulation 1 (Ern1) and Leu-rich Repeat Receptor-Like Kinase (RinRK1) are dispensable for intercellular infection in contrast to intracellular infection. Other symbiotic genes, including nod factor receptor 5 (NFR5), symbiosis receptor-like kinase (SymRK), Ca2+/calmodulin dependent kinase (CCaMK), exopolysaccharide receptor 3 (Epr3), Cyclops, nodule inception (Nin), nodulation signaling pathway 1 (Nsp1), nodulation signaling pathway 2 (Nsp2), cystathionine-β-synthase (Cbs), and Vapyrin are equally important for both entry modes. Comparative RNAseq analysis of roots inoculated with IRBG74 revealed a distinctive transcriptome response compared with intracellular colonization. In particular, several cytokinin-related genes were differentially regulated. Corroborating this observation, cyp735A and ipt4 cytokinin biosynthesis mutants were significantly affected in their nodulation with IRBG74, whereas lhk1 cytokinin receptor mutants formed no nodules. These results indicate a differential requirement for cytokinin signaling during intercellular rhizobial entry and highlight distinct modalities of inter- and intracellular infection mechanisms in L. japonicus., (© The Author(s) 2020. Published by Oxford University Press on behalf of American Society of Plant Biologists.)

The exocyst is a key factor in vesicle transport and is involved in cell secretion, cell growth, cell division and other cytological processes in eukaryotes. EXO70 is the key exocyst subunit. We obtained a gene, SHORT-ROOT 1 (SR1), through map-based cloning and genetic complementation. SR1 is a conserved protein with an EXO70 domain in plants. SR1 mutation affected the whole root-development process: producing shorter radicles, adventitious roots and lateral roots, and demonstrating abnormal xylem development, resulting in dwarfing and reduced water potential and moisture content. SR1 was largely expressed in the roots, but only in developing root meristems and tracheary elements. The shortness of the sr1 mutant roots was caused by the presence of fewer meristem cells. The in situ histone H4 expression patterns confirmed that cell proliferation during root development was impaired. Tracheary element dysplasia was caused by marked decreases in the inner diameters of and distances between the perforations of adjacent tracheary elements. The membrane transport of sr1 mutants was blocked, affecting cell division in the root apical region and the development of root tracheary elements. The study of SR1 will deepen our understanding of the function of EXO70 genes in Oryza sativa (rice) and guide future studies on the molecular mechanisms involved in plant root development., (© 2020 Society for Experimental Biology and John Wiley & Sons Ltd.)

Rose (Rosa chinensis) is the most important ornamental crops worldwide. However, the physiological and molecular mechanism of rose under drought stress remains elusive. In this study, we analyzed the changes of photosynthetic and phytohormone levels in the leaves and roots of rose seedlings grown under control (no drought), mild drought (MD) and severe drought stress. The total chlorophyll content and water use efficiency were significantly enhanced under MD in rose leaves. In addition, the concentration of ABA was higher in the leaves compared to the roots, whereas the roots accumulated more IAA, methylindole-3-acetic acid and indole-3-propionic acid. We also constructed the first full-length transcriptome for rose, and identified 96,201,862 full-length reads of average length 1,149 bp that included 65,789 novel transcripts. A total of 3,657 and 4,341 differentially expressed genes (DEGs) were identified in rose leaves and roots respectively. KEGG pathway analysis showed enrichment of plant hormone, signal transduction and photosynthesis are among the DEGs. 42,544 alternatively spliced isoforms were also identified, and alternative 3' splice site was the major alternative splicing (AS) event among the DEGs. Variations in the AS patterns of three genes between leaves and roots indicated the possibility of tissue-specific posttranscriptional regulation in response to drought stress. Furthermore, 2,410 novel long non-coding RNAs were detected that may participate in regulating the drought-induced DEGs. Our findings identified previously unknown splice sites and new genes in the rose transcriptome, and elucidated the drought stress-responsive genes as well as their intricate regulatory networks., (� The Author(s) 2020. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Circadian clock operates autonomously in each cell and drives the approximately 24-h rhythm in individual tissues and organs. It is known that the evening complex (EC) components and GI are required for ambient temperature perception and thermomorphogenesis in higher plants. Our previous study found that PRR9 and 7 are required for the lengthened period of the circadian rhythm in roots, and they are responsible for the temperature overcompensation in shoots. However, the molecular mechanism of the circadian clock, especially in different tissues, in response to temperature oscillations remains largely unknown. Here, we studied the transcript levels of EC genes and GI of the prr7 prr9 mutant shoots and roots in response to 22°C or 28°C, respectively. The results showed that PRR9 , 7 in roots inhibited the transcripts accumulation of ELF3, ELF4 , and LUX at 28°C. In addition, loss-of-function of both PRR9 and 7 caused an increase in GI expression at 22°C, but warm temperature of 28°C limited the negative effect of PRR9 , 7 on GI in roots. Our findings proposed a temperature-dependent molecular basis for root-specific circadian clock and indicated the critical role for PRR9 , 7 in negatively regulating ELF3, ELF4, LUX , and GI in the circadian gating of thermoresponse.

Environmental pH is a critical parameter for innumerable chemical reactions, myriad biological processes and all forms of life. The mechanisms that underlie the perception of external pH (pH e ) have been elucidated in detail for bacteria, fungi and mammalian cells; however, little information is available on whether and, if so, how pH e is perceived by plants. This is particularly surprising since hydrogen ion activity of the substrate is of paramount significance for plants, governing the availability of mineral nutrients, the structure of the soil microbiome and the composition of natural plant communities. Rapid changes in soil pH require constant readjustment of nutrient acquisition strategies, which is associated with dynamic alterations in gene expression. Referring to observations made in diverse experimental set-ups that unambiguously show that pH e per se affects gene expression, we hypothesize that sensing of pH e in plants is mandatory to prioritize responses to various simultaneously received environmental cues.

Spliceosomes are large complexes regulating pre-mRNA processing in eukaryotes. Arabidopsis JANUS encodes a putative subunit of spliceosome`. We recently demonstrated that JANUS plays an essential role during early embryogenesis and root meristem development. Instead of mediating pre-mRNA splicing as a subunit of spliceosome, JANUS regulates the transcription of key genes by recruiting RNA Polymerase II (Pol II). Here, we summarize our latest findings and provide insights into the regulation of JANUS during Arabidopsis development.

Roots grow asymmetrically, sometimes helically, around their growth direction likely to facilitate environmental sensing. We recently demonstrated that nitrate deficiency induces root coiling on horizontal surface through nitrate transporter/sensor NRT1.1 and PIN2- and AUX-mediated polar auxin transport. Here, we show that nitrate deficiency or NRT1.1 loss-of-function induces differential distribution of PIN2 between the future concave and concave sides in root epidermal cells. Treatment with pharmacological drugs suggests that enhanced endocytosis at the future convex side leads to reduced plasma membrane (PM) association of PIN2. A reduction of PIN2 at the PM would maintain a low auxin response to further enhance endocytosis at the convex side, leading to root coiling.

Earlier, we reported that Arabidopsis young lateral roots (LR) exhibited improved lethal salinity tolerance as compared with the primary root (PR). We have shown that cell death processes which take place in the PR during salt stress are postponed in the LR. Still, very little is known about the regulation of cell survival mechanisms in the LR during salinity stress. Here we used transgenic Arabidopsis plants expressing Cyclin B1;1:GUS, to further study the responses to salinity in the PR and LR positions. We found strong Cyclin B1;1:GUS activity in young budding LR of salt stressed and stress recovered plants. The Cyclin B1;1:GUS activity dropped significantly in long LR and was almost completely abolished in the PR. Our data provides another line of evidence that position-dependent response occurs in Arabidopsis roots during lethal salinity. The possible roles Cyclin B1;1 plays in the young LR during the response to lethal salinity are discussed.

Salt stress has negative impact on plant development and growth. Jasmonic acid (JA), a phytohormone, has been shown to involve in salt-induced inhibition of primary root growth. The Arabidopsis Glucosinolate transporter1 (GTR1/NPF2.10) is characterized as a JA-Ile, a bioactive form of JA, transporter. However, whether GTR1 participates in salt responses is not clear. In this study, we confirmed that GTR1 is induced by both JA and salinity. Salt-induced JA signaling is affected in gtr1 mutant. The JA responsive genes, JAZ1, JAZ5, MYC2, LOX3, are down-regulated in gtr1 mutant. Phenotypic analyses showed that the salinity-induced lateral root growth inhibition is enhanced in gtr1 mutant, suggesting that GTR1 plays a positive role in lateral root development under salt stress. Interestingly, the expression of a Na + transporter, HKT1, is upregulated in gtr1. Since HKT1 is a negative regulator for lateral root development under salt stress, we proposed that GTR1 alleviates the repression of lateral root development by salt stress by mediating JA signaling and repressing HKT1 expression. This study demonstrates that GTR1 is the molecular link between salt stress, JA signaling, and lateral root development., Competing Interests: Declaration of Competing Interest The authors have no conflicts of interest to declare., (Copyright © 2020 Elsevier B.V. All rights reserved.)

In our recent publication, we have shown that a member of the Laccase family, LACCASE2 ( LAC2) acted as a negative regulator of lignin deposition in the root xylem tissue of Arabidopsis thaliana. LAC2 messenger RNA (mRNA) level was post-transcriptionally regulated by microRNA 397b, which showed increased expression under water and phosphate deficiency, resulting in the downregulation of LAC2 expression In this report, we have investigated root growth and lignin deposition in an economically important legume crop chickpea ( Cicer arietinum L.) in response to natural drought in soil-grown condition. In contrast to the growth retardation of Arabidopsis root in mannitol-supplemented medium, chickpea root showed an increase in length in low soil moisture condition. Lignin estimation in the primary root showed an increase in lignin content, which was substantiated by staining of root xylem. Drought treatment enhanced the expression of four out of six LAC genes tested, while the expression of two was downregulated. Our preliminary study indicateed a molecular mechanism of lignin deposition in chickpea root xylem during drought.

The interaction of C-TERMINALLY ENCODED PEPTIDES (CEPs) with CEP RECEPTOR1 (CEPR1) controls root growth and development, as well as nitrate uptake, but has no known role in determining yield. We used physiological, microscopic, molecular, and grafting approaches to demonstrate a reproductive tissue-specific role for CEPR1 in controlling yield and seed size. Independent Arabidopsis ( Arabidopsis thaliana ) cepr1 null mutants showed disproportionately large reductions in yield and seed size relative to their decreased vegetative growth. These yield defects correlated with compromised reproductive development predominantly in female tissues, as well as chlorosis, and the accumulation of anthocyanins in cepr1 reproductive tissues. The thinning of competing reproductive organs to improve source-to-sink ratios in cepr1 , along with reciprocal bolt-grafting experiments, demonstrated that CEPR1 acts locally in the reproductive bolt to control yield and seed size. CEPR1 is expressed throughout the vasculature of reproductive organs, including in the chalazal seed coat, but not in other seed tissues. This expression pattern implies that CEPR1 controls yield and seed size from the maternal tissue. The complementation of cepr1 mutants with transgenic CEPR1 rescued the yield and other phenotypes. Transcriptional analyses of cepr1 bolts showed alterations in the expression levels of several genes of the CEP-CEPR1 and nitrogen homeostasis pathways. This transcriptional profile was consistent with cepr1 bolts being nitrogen deficient and with a reproductive tissue-specific function for CEP-CEPR1 signaling. The results reveal a local role for CEPR1 in the maternal reproductive tissue in determining seed size and yield, likely via the control of nitrogen delivery to the reproductive sinks., (© 2020 American Society of Plant Biologists. All Rights Reserved.)

Calcium (Ca 2+ ) as a universal signal molecule plays pivotal roles in plant growth and development. It regulates root morphogenesis mainly through mediating phytohormone and stress signalings or affecting these signalings. In recent years, much progress has been made in understanding the roles of Ca 2+ in primary root development. Here, we summarize recent advances in the functions and mechanisms of Ca 2+ in modulating primary root growth in plants under normal and stressful conditions.

Root systems are dynamic and adaptable organs that play critical roles in plant development. However, how roots grow and accumulate biomass during plant life cycle and in relation to shoot growth phenology remains understudied. A comprehensive time-dependent root morphological analysis integrated with molecular signatures is then required to advance our understanding of root growth and development. Here we studied Brachypodium distachyon rooting process by monitoring root morphology, biomass production, and C/N ratios during developmental stages. To provide insight into gene regulation that accompanies root growth, we generated comprehensive transcript profiles of Brachypodium whole-root system at four developmental stages. Our data analysis revealed that multiple biological processes including trehalose metabolism and various families of transcription factors (TFs) were differentially expressed in root system during plant development. In particular, the AUX/IAA, ERFs, WRKY, NAC, and MADS TF family members were upregulated as plant entered the booting/heading stage, while ARFs and GRFs were downregulated suggesting these TF families as important factors involved in specific phases of rooting, and possibly in regulation of transition to plant reproductive stages. We identified several Brachypodium candidate root biomass-promoting genes and cis-regulatory elements for further functional validations and root growth improvements in grasses.

The plasma membrane intrinsic protein PIP2;5 is the most highly expressed aquaporin in maize ( Zea mays ) roots. Here, we investigated how deregulation of PIP2;5 expression affects water relations and growth using maize overexpression (OE; B104 inbred) or knockout (KO; W22 inbred) lines. The hydraulic conductivity of the cortex cells of roots grown hydroponically was higher in PIP2;5 OE and lower in pip2;5 KO lines compared with the corresponding wild-type plants. While whole-root conductivity decreased in the KO lines compared to the wild type, no difference was observed in OE plants. This paradox was interpreted using the MECHA hydraulic model, which computes the radial flow of water within root sections. The model hints that the plasma membrane permeability of the cells is not radially uniform but that PIP2;5 may be saturated in cell layers with apoplastic barriers, i.e. the endodermis and exodermis, suggesting the presence of posttranslational mechanisms controlling the abundance of PIP in the plasma membrane in these cells. At the leaf level, where the PIP2;5 gene is weakly expressed in wild-type plants, the hydraulic conductance was higher in the PIP2;5 OE lines compared with the wild-type plants, whereas no difference was observed in the pip2;5 KO lines. The temporal trend of leaf elongation rate, used as a proxy for that of xylem water potential, was faster in PIP2;5 OE plants upon mild stress, but not in well-watered conditions, demonstrating that PIP2;5 may play a beneficial role in plant growth under specific conditions., (© 2020 American Society of Plant Biologists. All Rights Reserved.)

Thompson Seedless, a commonly grown table grape variety, is sensitive to salinity when grown on its own roots, and therefore, it is frequently grafted onto salinity-tolerant wild grapevine rootstocks. Rising soil salinity is a growing concern in irrigated agricultural systems. The accumulation of salts near the root zone severely hampers plant growth, leading to a decrease in the productive lifespan of grapevine and causing heavy yield losses to the farmer. In the present study, we investigated the differences in response to salinity between own-rooted Thompson Seedless (TSOR) and 110R-grafted Thompson Seedless (TS110R) grapevines, wherein 110R is reported to be a salt-tolerant rootstock. The grapevines were subjected to salt stress by treating them with a 150 mM NaCl solution. The stress-induced changes in protein abundance were investigated using a label-free shotgun proteomics approach at three time-points viz. 6 h, 48 h, and 7 days of salt treatment. A total of 2793 proteins were identified, of which 246 were differentially abundant at various time-points in TSOR and TS110R vines. The abundance of proteins involved in several biological processes such as photosynthesis, amino acid metabolism, translation, chlorophyll biosynthesis, and generation of precursor metabolites was significantly affected by salt stress in both the vines but at different stages of stress. The results revealed that TSOR vines responded fervently to salt stress, while TS110R vines adopted a preventive approach. The findings of this study add to the knowledge of salinity response in woody and grafted plants and hence open the scope for further studies on salt stress-specific differences induced by grafting.

Potato (Solanum tuberosum) mutant (ST) lacking one isoform of manganese-stabilizing protein (MSPI) of photosystem II exhibited besides spontaneous tuberization also growth changes with strongly impaired root system development. Previous studies revealed marked changes in carbohydrate levels and allocation within ST plant body. To verify causal relationship between changed carbohydrate balance and root growth restriction we engaged dark grown sucrose-supplied root organ-cultures of ST plants to exclude/confirm shoot effects. Unexpectedly, in ST root cultures we observed large alterations in growth and architecture as well as saccharide status similar to those found in the intact plant roots. The gene expression analysis, however, proved PsbO1 transcript (coding MSPI protein) neither in ST nor in WT root-organ cultures. Therefore, the results point to indirect effects of PsbO1 allele absence connected possibly with some epigenetic modulations., Competing Interests: Declaration of Competing Interest None., (Copyright © 2019 Elsevier GmbH. All rights reserved.)

The mechanisms involved in the regulation of gene expression in response to phosphate (Pi) deficiency have been extensively studied, but their chromatin-level regulation remains poorly understood. We examined the role of histone acetylation in response to Pi deficiency by using the histone deacetylase complex1 (hdc1) mutant. Genes involved in root system architecture (RSA) remodeling were analyzed by quantitative real-time polymerase chain reaction (qPCR) and chromatin immunoprecipitation qPCR. We demonstrate that histone H3 acetylation increased under Pi deficiency, and the hdc1 mutant was hypersensitive to Pi deficiency, with primary root growth inhibition and increases in root hair number. Concomitantly, Pi deficiency repressed HDC1 protein abundances. Under Pi deficiency, hdc1 accumulated higher concentrations of Fe 3+ in the root tips and had higher expression of genes involved in RSA remodeling, such as ALUMINUM-ACTIVATED MALATE TRANSPORTER1 (ALMT1), LOW PHOSPHATE ROOT1 (LPR1), and LPR2 compared with wild-type plants. Furthermore, Pi deficiency enriched the histone H3 acetylation of ALMT1 and LPR1. Finally, genetic evidence showed that LPR1/2 was epistatic to HDC1 in regulating RSA remodeling. Our results suggest a chromatin-level control of Pi starvation responses in which HDC1-mediated histone H3 deacetylation represses the transcriptional activation of genes involved in RSA remodeling in Arabidopsis., (© 2019 The Authors. New Phytologist © 2019 New Phytologist Trust.)

OsCASP1 (Casparian strip domain protein 1) was recently identified to function in Casparian strip (CS) formation at the endodermal cells in rice roots, which was required for selective mineral uptake in rice. Here, we further investigate the functional form of OsCASP1 in vivo . Expression analysis shows that OsCASP1, OsCASP2, OsCASP3 , and OsCASP5 were expressed in roots apart from OsCASP4 . A yeast two-hybrid (Y2H) assay revealed that OsCASP1 can interact with itself and OsCASP2, but not with OsCASP3 and OsCASP5. These interactions of OsCASP1 with itself and OsCASP2 at the plasma membrane were confirmed using bimolecular fluorescence complementation (BiFC) in rice protoplasts. These results indicated that OsCASP1 can form complexes with itself and OsCASP2 in rice roots.

The heterogeneous distribution of soil salinity across the rhizosphere can moderate salt injury and improve sorghum growth. However, the essential molecular mechanisms used by sorghum to adapt to such environmental conditions remain uncharacterized. The present study evaluated physiological parameters such as the photosynthetic rate, antioxidative enzyme activities, leaf Na+ and K+ contents, and osmolyte contents and investigated gene expression patterns via RNA sequencing (RNA-seq) analysis under various conditions of nonuniformly distributed salt. Totals of 5691 and 2047 differentially expressed genes (DEGs) in the leaves and roots, respectively, were identified by RNA-seq under nonuniform (NaCl-free and 200 mmol·L-1 NaCl) and uniform (100 mmol·L-1 and 100 mmol·L-1 NaCl) salinity conditions. The expression of genes related to photosynthesis, Na+ compartmentalization, phytohormone metabolism, antioxidative enzymes, and transcription factors (TFs) was enhanced in leaves under nonuniform salinity stress compared with uniform salinity stress. Similarly, the expression of the majority of aquaporins and essential mineral transporters was upregulated in the NaCl-free root side in the nonuniform salinity treatment, whereas abscisic acid (ABA)-related and salt stress-responsive TF transcripts were more abundant in the high-saline root side in the nonuniform salinity treatment. In contrast, the expression of the DEGs identified in the nonuniform salinity treatment remained virtually unaffected and was even downregulated in the uniform salinity treatment. The transcriptome findings might be supportive of the increased photosynthetic rate, reduced Na+ levels, increased antioxidative capability in the leaves and, consequently, the growth recovery of sorghum under nonuniform salinity stress as well as the inhibited sorghum growth under uniform salinity conditions. The increased expression of salt resistance genes activated in response to the nonuniform salinity distribution implied that the cross-talk between the nonsaline and high-saline sides of the roots exposed to nonuniform salt stress is potentially regulated., Competing Interests: The authors have declared that no competing interests exist.

Anchor roots (ANRs) arise at the root-shoot junction and are the least investigated type of Arabidopsis root. Here, we show that ANRs originate from pericycle cells in an auxin-dependent manner and a carotenogenic signal to emerge. By screening known and assumed carotenoid derivatives, we identified anchorene, a presumed carotenoid-derived dialdehyde (diapocarotenoid), as the specific signal needed for ANR formation. We demonstrate that anchorene is an Arabidopsis metabolite and that its exogenous application rescues the ANR phenotype in carotenoid-deficient plants and promotes the growth of normal seedlings. Nitrogen deficiency resulted in enhanced anchorene content and an increased number of ANRs, suggesting a role of this nutrient in determining anchorene content and ANR formation. Transcriptome analysis and treatment of auxin reporter lines indicate that anchorene triggers ANR formation by modulating auxin homeostasis. Together, our work reveals a growth regulator with potential application to agriculture and a new carotenoid-derived signaling molecule., (Copyright © 2019 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. Distributed under a Creative Commons Attribution NonCommercial License 4.0 (CC BY-NC).)

The Arabidopsis ( Arabidopsis thaliana ) root epidermis exhibits a position-dependent pattern of root-hair and nonhair cell types. A highly orchestrated network of gene regulatory interactions, including the R2R3-type MYB transcription factor WEREWOLF (WER), is responsible for generating this cell pattern during root development. In this study, we identified a novel wer mutant from a genetic enhancer screen, designated wer-4 , that exhibits an abnormal pattern of root-hair and nonhair cells. We established that wer-4 bears a single-residue substitution (D105N) in the DNA-binding R3 MYB repeat of WER, which differentially affects the transcription of WER target genes, including GLABRA2 , CAPRICE , TRIPTYCHON , and ENHANCER OF TRY AND CPC1 This modulation of the gene regulatory network leads to altered levels and distributions of cell fate regulators in the differentiating epidermal cells that ultimately generate the abnormal cell-type pattern. We also created several WER variants with substitutions at the Asp-105 position, and these exhibited a variety of gene expression and cell-type pattern alterations, further supporting the critical role of this residue. These findings provide insight into WER protein function and its importance in generating the proper balance of downstream transcriptional factors in the gene regulatory network that establishes root epidermal cell fate., (© 2019 American Society of Plant Biologists. All Rights Reserved.)

Zinc (Zn) deficiency is a widespread agricultural problem in arable soils of the whole world. However, the molecular mechanisms underlying Zn-deficiency response are largely unknown. Here, we analyzed the transcriptomic profilings of soybean leaves and roots in response to Zn deficiency through Illumina's high-throughput RNA sequencing in order to understand the molecular basis of Zn-deficiency response in the plants. A total of 614 and 1011 gene loci were found to be differentially expressed in leaves and roots, respectively, and 88 loci were commonly found in both leaves and roots. Twelve differentially expressed genes (DEGs) were randomly selected for validation by quantitative reverse transcription polymerase chain reaction, and their fold changes were similar to those of RNA-seq. Gene ontology enrichment analysis showed that ion transport, nicotianamine (NA) biosynthetic process and queuosine biosynthetic process were enriched in the upregulated genes, while oxidation-reduction process and defense response were enriched in the downregulated genes. Among the DEGs, 20 DEGs are potentially involved in Zn homeostasis, including seven ZRT, IRT-related protein (ZIP) transporter genes, three NA synthase genes, and seven metallothionein genes; 40 DEGs are possibly involved in diverse hormonal signals such as auxin, cytokinin, ethylene and gibberellin; nine DEGs are putatively involved in calcium signaling; 85 DEGs are putative transcription factor genes. Nine DEGs were found to contain zinc-deficiency-response element in their promoter regions. These results could provide comprehensive insights into the soybean response to Zn deficiency and will be helpful for further elucidation of the molecular mechanisms of Zn-deficiency response and Zn-deficiency tolerance in plants., (© 2018 Scandinavian Plant Physiology Society.)

The phloem, located within the vascular system, is critical for delivery of nutrients and signaling molecules throughout the plant body. Although the morphological process and several factors regulating phloem differentiation have been reported, the molecular mechanism underlying its initiation remains largely unknown. Here, we report that the small peptide-coding gene, CLAVATA 3 (CLV3)/EMBEYO SURROUNDING REGION 25 (CLE25), the expression of which begins in provascular initial cells of 64-cell-staged embryos, and continues in sieve element-procambium stem cells and phloem lineage cells, during post-embryonic root development, facilitates phloem initiation in Arabidopsis. Knockout of CLE25 led to delayed protophloem formation, and in situ expression of an antagonistic CLE25 G6T peptide compromised the fate-determining periclinal division of the sieve element precursor cell and the continuity of the phloem in roots. In stems of CLE25 G6T plants the phloem formation was also compromised, and procambial cells were over-accumulated. Genetic and biochemical analyses indicated that a complex, consisting of the CLE-RESISTANT RECEPTOR KINASE (CLERK) leucine-rich repeat (LRR) receptor kinase and the CLV2 LRR receptor-like protein, is involved in perceiving the CLE25 peptide. Similar to CLE25, CLERK was also expressed during early embryogenesis. Taken together, our findings suggest that CLE25 regulates phloem initiation in Arabidopsis through a CLERK-CLV2 receptor complex., (© 2019 Institute of Botany, Chinese Academy of Sciences.)

The SPORULATION 11 (SPO11) proteins are among eukaryotic the topoisomerase VIA (Topo VIA) homologs involved in modulating various important biological processes, such as growth, development and stress response via endoreduplication in plants, but the underlying mechanism response to stress remains largely unknown under salt treatment. Here, we attempted to characterize a homolog of TOP VIA in upland cotton (Gossypium hirsutum L.), designated as GhSPO11-3. The silencing of GhSPO11-3 in cotton plants resulted in a dwarf phenotype with a failure of cell endoreduplication and a phase shift in the ploidy levels. The GhSPO11-3-silenced plants also showed substantial changes including accumulated malondialdehyde, significantly reduced chlorophyll and proline contents and decreased antioxidative enzyme activity after salt treatment. In addition, transgenic Arabidopsis lines overexpressing GhSPO11-3 accelerated both leaf and root growth with cell expansion and endopolyploidy. Both leaf stomatal density and aperture were markedly decreased, and the transgenic Arabidopsis lines were more tolerant with expression of stress-responsive genes under salinity stress. Furthermore, consistent with the reduced reactive oxygen species (ROS), the expression of ROS scavenging-related genes was largely reinforced, and antioxidant enzyme activities were accordingly significantly enhanced in transgenic Arabidopsis lines under salt stress. In general, these results indicated that GhSPO11-3 likely respond to salt stress by positively regulating root growth, stomatal response, ROS production and the expression of stress-related genes to cope with adverse conditions in plants., (© 2018 Scandinavian Plant Physiology Society.)

Adventitious root (AR) is a kind of later root, which derives from stems and leaf petioles of plants. Many different kinds of small signaling molecules can transmit information between cells of multicellular organisms. It has been found that small molecules can be involved in many growth and development processes of plants, including stomatal movement, flowering, fruit ripening and developing, and AR formation. Therefore, this review focuses on discussing the functions and mechanisms of small signaling molecules in the adventitious rooting process. These compounds, such as nitric oxide (NO), hydrogen gas (H 2 ), hydrogen sulfide (H 2 S), carbon monoxide (CO), methane (CH 4 ), ethylene (ETH), and hydrogen peroxide (H 2 O 2 ), can be involved in the induction of AR formation or development. This review also sums the crosstalk between these compounds. Besides, those signaling molecules can regulate the expressions of some genes during AR development, including cell division genes, auxin-related genes, and adventitious rooting-related genes. We conclude that these small-molecule compounds enhance adventitious rooting by regulating antioxidant, water balance, and photosynthetic systems as well as affecting transportation and distribution of auxin, and these compounds further conduct positive effects on horticultural plants under environmental stresses. Hence, the effect of these molecules in plant AR formation and development is definitely a hot issue to explore in the horticultural study now and in the future.

Lateral root (LR) proliferation is a major determinant of soil nutrient uptake. How resource allocation controls the extent of LR growth remains unresolved. We used genetic, physiological, transcriptomic, and grafting approaches to define a role for C-TERMINALLY ENCODED PEPTIDE RECEPTOR 1 (CEPR1) in controlling sucrose-dependent LR growth. CEPR1 inhibited LR growth in response to applied sucrose, other metabolizable sugars, and elevated light intensity. Pathways through CEPR1 restricted LR growth by reducing LR meristem size and the length of mature LR cells. RNA-sequencing of wild-type (WT) and cepr1-1 roots with or without sucrose treatment revealed an intersection of CEP-CEPR1 signalling with the sucrose transcriptional response. Sucrose up-regulated several CEP genes, supporting a specific role for CEP-CEPR1 in the response to sucrose. Moreover, genes with basally perturbed expression in cepr1-1 overlap with WT sucrose-responsive genes significantly. We found that exogenous CEP inhibited LR growth via CEPR1 by reducing LR meristem size and mature cell length. This result is consistent with CEP-CEPR1 acting to curtail the extent of sucrose-dependent LR growth. Reciprocal grafting indicates that LR growth inhibition requires CEPR1 in both the roots and shoots. Our results reveal a new role for CEP-CEPR1 signalling in controlling LR growth in response to sucrose., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

The plant body plan and primary organs are established during embryogenesis. However, in contrast to animals, plants have the ability to generate new organs throughout their whole life. These give them an extraordinary developmental plasticity to modulate their size and architecture according to environmental constraints and opportunities. How this plasticity is regulated at the whole-organism level is elusive. Here we provide evidence for a role for translationally controlled tumour protein (TCTP) in regulating the iterative formation of lateral roots in Arabidopsis. AtTCTP1 modulates root system architecture through a dual function: as a general constitutive growth promoter enhancing root elongation and as a systemic signalling agent via mobility in the vasculature. AtTCTP1 encodes mRNAs with long-distance mobility between the shoot and roots. Mobile shoot-derived TCTP1 gene products act specifically to enhance the frequency of lateral root initiation and emergence sites along the primary root pericycle, while root elongation is controlled by local constitutive TCTP1 expression and scion size. These findings uncover a novel type for an integrative signal in the control of lateral root initiation and the compromise for roots between branching more profusely or elongating further. They also provide the first evidence in plants of an extracellular function of the vital, highly expressed ubiquitous TCTP1., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Society for Experimental Biology.)

Brassinosteroid (BR) plays an important role in plant development and biotic and abiotic stress tolerance, but its specific function remains largely unknown in wheat (Triticum aestivum L.), preventing its utilization in this important crop. In this study, the function of BR and its underlying cytological role in wheat root development were comprehensively investigated. Our findings demonstrated that BR has a conserved function in regulating root length in wheat, and novel roles in regulating lateral root emergence and root diameter were uncovered. Analyses of BR homologous gene composition and evolutionary divergence demonstrated that the genetic framework of the wheat BR pathway was close to that of rice, but contained highly redundant homologous copies of genes from the subgenome A, B and D. These homologous copies showed active expression and shared a conserved BR response. The expression of wheat DWF4 and glycogen synthase kinase (GSK) genes in Arabidopsis confirmed that multiple homologous copies maintained their conserved function in regulating root development, highlighting their redundant status and indicating that a special challenge exists in wheat gene modification to deal with this high redundancy. However, our results suggested that the hypermorphic effect of T. aestivum GSK (TaGSK) genes with point mutations may be an effective approach to overcome this redundancy in the manipulation of BR signaling in wheat. Our study provides fundamental data uncovering the function of BR in wheat root development, the underlying genetic basis and a possible strategy to manipulate BR signaling in hexaploid wheat., (� The Author(s) 2019. Published by Oxford University Press on behalf of Japanese Society of Plant Physiologists. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Key Message: Overexpression of SlMBP9 reduced auxin biosynthesis and transport, and negatively regulated lateral root formation and apical dominance. MADS-box transcription factors play a critical role in plant development. In this study, we describe SlMBP9, a novel MADS-box gene that is expressed in the roots of tomato plants. Tomato lines that over- or under-expressed SlMBP9 were generated using a transgenic approach. The number of lateral roots (LRs) were reduced in SlMBP9-overexpressing lines but slightly increased in SlMBP9-silenced lines. A physiological index revealed that the auxin content significantly decreased in the root maturation zone of the overexpression lines. In addition, gene expression analysis revealed that the expression of the polar auxin transporter genes PIN1 and ABCB19/MDR1 and genes involved in auxin biosynthesis was downregulated in the stems of overexpression lines, which is consistent with the reduced accumulation of auxin in the root maturation zone. Exogenous indole-3-acetic acid (auximone) rescued the lateral root phenotypes of the SlMBP9-overexpressing lines. Overexpression of SlMBP9 resulted in dwarf plants, enhanced lateral buds and reduced the gibberellin content in the stems. Together, these results suggest that SlMBP9 plays a negative role in the process of auxin biosynthesis and transport.

Desert plants have developed mechanisms for adapting to hostile desert conditions, yet these mechanisms remain poorly understood. Here, we describe two unique modes used by desert date palms ( Phoenix dactylifera ) to protect their meristematic tissues during early organogenesis. We used x-ray micro-computed tomography combined with high-resolution tissue imaging to reveal that, after germination, development of the embryo pauses while it remains inside a dividing and growing cotyledonary petiole. Transcriptomic and hormone analyses show that this developmental arrest is associated with the low expression of development-related genes and accumulation of hormones that promote dormancy and confer resistance to stress. Furthermore, organ-specific cell-type mapping demonstrates that organogenesis occurs inside the cotyledonary petiole, with identifiable root and shoot meristems and their respective stem cells. The plant body emerges from the surrounding tissues with developed leaves and a complex root system that maximizes efficient nutrient and water uptake. We further show that, similar to its role in Arabidopsis ( Arabidopsis thaliana ), the SHORT-ROOT homolog from date palms functions in maintaining stem cell activity and promoting formative divisions in the root ground tissue. Our findings provide insight into developmental programs that confer adaptive advantages in desert plants that thrive in hostile habitats., (© 2019 The author(s).)

Plants respond to herbivore or pathogen attacks by activating specific defense programs that include the production of bioactive specialized metabolites to eliminate or deter the attackers. Volatiles play an important role in the interaction of a plant with its environment. Through transcript profiling of jasmonate-elicited Medicago truncatula cells, we identified Emission of Methyl Anthranilate (EMA) 1, a MYB transcription factor that is involved in the emission of the volatile compound methyl anthranilate when expressed in M. truncatula hairy roots, giving them a fruity scent. RNA sequencing (RNA-Seq) analysis of the fragrant roots revealed the upregulation of a methyltransferase that was subsequently characterized to catalyze the O-methylation of anthranilic acid and was hence named M. truncatula anthranilic acid methyl transferase (MtAAMT) 1. Given that direct activation of the MtAAMT1 promoter by EMA1 could not be unambiguously demonstrated, we further probed the RNA-Seq data and identified the repressor protein M. truncatula plant AT-rich sequence and zinc-binding (MtPLATZ) 1. Emission of Methyl Anthranilate 1 binds a tandem repeat of the ACCTAAC motif in the MtPLATZ1 promoter to transactivate gene expression. Overexpression of MtPLATZ1 in transgenic M. truncatula hairy roots led to transcriptional silencing of EMA1, indicating that MtPLATZ1 may be part of a negative feedback loop to control the expression of EMA1. Finally, application of exogenous methyl anthranilate boosted EMA1 and MtAAMT1 expression dramatically, thus also revealing a positive amplification loop. Such positive and negative feedback loops seem to be the norm rather than the exception in the regulation of plant specialized metabolism., (© 2019 The Authors The Plant Journal © 2019 John Wiley & Sons Ltd.)

The major root system of cereals consists of crown roots (or adventitious roots), which are important for anchoring plants in the soil and for water and nutrient uptake. However, the molecular basis of crown root formation is largely unknown. Here, we isolated a rice ( Oryza sativa ) mutant with fewer crown roots, named lower crown root number1 ( lcrn1 ). Map-based cloning revealed that lcrn1 is caused by a mutation of a putative transcription factor-coding gene, O. sativa SQUAMOSA PROMOTER BINDING PROTEIN-LIKE3 ( OsSPL3 ). We demonstrate that the point mutation in lcrn1 perturbs the O. sativa microRNA156 (OsmiR156)-directed cleavage of OsSPL3 transcripts, resulting in the mutant phenotype. Chromatin immunoprecipitation sequencing assays of OsSPL3 binding sites and RNA sequencing of differentially expressed transcripts in lcrn1 further identified potential direct targets of OsSPL3 in basal nodes, including a MADS-box transcription factor, OsMADS50. OsMADS50- overexpressing plants produced fewer crown roots, phenocopying lcrn1 , while knocking out OsMADS50 in the lcrn1 background reversed this phenotype. We also show that OsSPL12 , another OsmiR156 target gene, regulates OsMADS50 and crown root development. Taken together, our findings suggest a novel regulatory pathway in which the OsmiR156- OsSPL3/OsSPL12 module directly activates OsMADS50 in the node to regulate crown root development in rice., (© 2019 American Society of Plant Biologists. All rights reserved.)

White lupin cluster roots are specialized brush-like root structures that are formed in some species under phosphorus (P)-deficient conditions. They intensely secrete protons and organic acid anions for solubilization and acquisition of sparingly soluble phosphates. Phytohormones and sucrose modulate cluster root number, but the molecular mechanisms of cluster root formation have been elusive. Here, a novel peptide phytohormone was identified that affects cluster root development. It belongs to the C-TERMINALLY-ENCODED PEPTIDE (CEP) family. Members of that family arrest root growth and modulate branching in model species. LaCEP1 was highly expressed in the pre-emergence zone of clusters. Over-expression of the gene encoding the LaCEP1 propeptide resulted in moderate inhibition of cluster root formation. The primary and lateral root lengths of lupin were little affected by the overexpression, but LaCEP1 reduced cluster rootlet and root hair elongation. Addition of a 15-mer core peptide derived from LaCEP1 similarly altered root morphology and modified cluster activity, suggesting that a core sequence of the propeptide is functionally sufficient. Stable overexpression in Arabidopsis confirmed the LaCEP1 function in root growth inhibition across species. Taken together, the root inhibitory effects of the LaCEP1 phytohormone suggest a role as of a regulatory module involved in cluster root development in white lupin., (© 2018 Scandinavian Plant Physiology Society.)

Root hairs play important roles for the acquisition of nutrients, microbe interaction and plant anchorage. In addition, root hairs provide an excellent model system to study cell patterning, differentiation and growth. Arabidopsis root hairs have been thoroughly studied to understand how plants regulate cell fate and growth in response to environmental signals. Accumulating evidence suggests that a multi-layered gene regulatory network is the molecular secret to enable the flexible and adequate response to multiple signals. In this review, we describe the key transcriptional regulators controlling cell fate and/or cell growth of root hairs. We also discuss how plants integrate phytohormonal and environmental signals, such as auxin, ethylene and phosphate availability, and modulate the level of these transcriptional regulators to tune root hair development.

Chelidonium majus is a traditional medicinal plant, which commonly known as a rich resource for the major benzylisoquinoline alkaloids (BIAs), including morphine, sanguinarine, and berberine. To understand the biosynthesis of C. majus BIAs, we performed de novo transcriptome sequencing of its leaf and root tissues using Illumina technology. Following comprehensive evaluation of de novo transcriptome assemblies produced with five programs including Trinity, Bridger, BinPacker, IDBA-tran, and Velvet/Oases using a series of k-mer sizes (from 25 to 91), BinPacker was found to produce the best assembly using a k-mer of 25. This study reports the results of differential gene expression (DGE), functional annotation, gene ontology (GO) analysis, classification of transcription factor (TF)s, and SSR and miRNA discovery. Our DGE analysis identified 6,028 transcripts that were up-regulated in the leaf, and 4,722 transcripts that were up-regulated in the root. Further investigations showed that most of the genes involved in the BIA biosynthetic pathway are significantly expressed in the root compared to the leaf. GO analysis showed that the predominant GO domain is "cellular component", while TF analysis found bHLH to be the most highly represented TF family. Our study further identified 10 SSRs, out of a total of 39,841, that showed linkage to five unigenes encoding enzymes in the BIA pathway, and 10 conserved miRNAs that were previously not detected in this plant. The comprehensive transcriptome information presented herein provides a foundation for further explorations on study of the molecular mechanisms of BIA synthesis in C. majus., Competing Interests: The authors have declared that no competing interests exist.

As sessile organisms, plants can modify their growth strategy in response to different temperatures, however very little is known about how roots growth responds to ambient temperature change. Here, we found that high temperature-induced root elongation is dependent on light intensity and the root growth of most TAA1 loss-of-function mutants is more sensitive to higher temperatures in Arabidopsis. TAA1 encodes a tryptophan aminotransferase which involved in the indole-3-pyruvic acid (IPA) pathway of indole-3-acetic acid (IAA) biosynthesis. The root elongation in ckrc1-1(one allele mutant of TAA1) is less sensitive to lower temperatures and more sensitive to higher temperatures than that of Col-0. By comparing the regulatory mechanisms of ckrc1-1 root growth at different temperatures (17 °C, 22 °C, and 27 °C), different interactions between signals (auxin and ethylene) and the effects of downstream genes were observed at different ambient temperatures in Arabidopsis. Lower temperature-enhanced ETR1-mediated ethylene signaling did not promote the expression of CKRC1, while higher temperature-enhanced signaling did. CKRC1 had an important role in the ACC inhibition of cell elongation at 22 °C and 27 °C but not at 17 °C. CKRC1-dependent auxin biosynthesis was critical for maintaining PIN1, PIN2, and AUX1 expression at lower temperatures. CKRC1, AUX1, and PIN2 regulated root elongation by affecting different regions of the root at different temperatures in Arabidopsis. Our experimental results suggested that changes in the in vivo signals at different temperatures were multi-layered in Arabidopsis., (Copyright © 2019 Elsevier B.V. All rights reserved.)

Plant responses to phosphate deprivation encompass a wide range of strategies, varying from altering root system architecture, entering symbiotic interactions to excreting root exudates for phosphorous release, and recycling of internal phosphate. These processes are tightly controlled by a complex network of proteins that are specifically upregulated upon phosphate starvation. Although the different effects of phosphate starvation have been intensely studied, the full extent of its contribution to altered root system architecture remains unclear. In this review, we focus on the effect of phosphate starvation on the developmental processes that shape the plant root system and their underlying molecular pathways., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Key Message: Pharmacological and molecular evidence reveals a novel role of methane (CH 4 ) gas in root organogenesis, the induction of lateral root (LR) formation, and this response might require hydrogen peroxide (H 2 O 2 ) synthesis. Although plants can produce CH 4 and release this to atmosphere, the beneficial role(s) of CH 4 are not fully elucidated. In this study, the fumigation with CH 4 not only increased NADPH oxidase activity and H 2 O 2 production, but also induced tomato lateral root primordial formation and thereafter LR development. However, exogenously applied argon and nitrogen failed to influence LR formation. Above responses triggered by CH 4 were sensitive to the removal of endogenous H 2 O 2 with dimethylthiourea (DMTU; a membrane-permeable scavenger of H 2 O 2 ), suggesting the hypothesis that CH 4 's effect on LR formation could be mediated by endogenous H 2 O 2 . Diphenylene iodonium (DPI) inhibition of the H 2 O 2 generating enzyme NADPH oxidase attenuated H 2 O 2 synthesis and impaired LR formation in response to CH 4 , confirming the requirement of NADPH oxidase-dependent H 2 O 2 . Meanwhile, the alterations of endogenous H 2 O 2 concentrations failed to influence CH 4 production in tomato seedlings. Molecular evidence revealed that CH 4 -induced SlCDKA1, SlCYCA2;1, and SlCYCA3;1 transcripts, and -decreased SlKRP2 mRNA were impaired by DMTU or DPI. Contrasting changes in LR formation-related miR390a and miR160 transcripts and their target genes, including SlARF4 and SlARF16, were observed. Together, our pharmacological and molecular evidence suggested the requirement of H 2 O 2 synthesis in CH 4 -triggered tomato LR formation, partially via the regulation of cell cycle regulatory genes, miRNA-, and tasiRNA-modulated gene expression.

Identification of root-specific promoters is a good method to drive root-specific gene expression for nutrient uptake. Constitutive over-expression of OsPT2 may have negative effects on the growth of rice seedlings under high Pi condition. Thus, characterization and utilization of root-specific promoters are critical for genetic breeding. Here, a root-specific promoter (Os03g01700) with a number of specific regulatory elements has been confirmed. Interestingly, cis-regulatory element S449 is significantly enriched in the -1475∼-2013 bp and -1077∼-1475 bp regions of Os03g01700 promoter. The activities of several deletion derivatives of Os03g01700 promoter were analyzed using both transient expression and genetic transformation system. The results showed that the root-specific cis-acting elements might be present in the -2013 bp～-1475 bp and -1077 bp～-561 bp regions of Os03g01700 promoter. To determine the actual effect of root-specific expression of OsPT2, a construction consisting of Os03g01700 promoter and OsPT2 CDS was used to transform rice. Under Pi-sufficient condition, there were a series of symptoms of phosphorus toxicity in the shoots of OsPT2 over-expressing (Ov-OsPT2) seedlings. Under Pi-deficient condition, more soluble Pi was accumulated in the shoots of Ov-OsPT2 seedlings than that in the wild type. Our data provide a candidate root-specific promoter in the breeding of rice with high phosphorus uptake variety., (Copyright © 2019. Published by Elsevier Masson SAS.)

In plants, the position of lateral roots (LRs) depends on initiation sites induced by auxin. The domain of high auxin response responsible for LR initiation stretches over several cells, but only a pair of pericycle cells (LR founder cells) will develop into LRs. In this work, we identified a signaling cascade controlling LR formation through lateral inhibition. It comprises a peptide hormone TARGET OF LBD SIXTEEN 2 (TOLS2), its receptor RLK7, and a downstream transcription factor PUCHI. TOLS2 is expressed at the LR founder cells and inhibits LR initiation. Time-lapse imaging of auxin-responsive DR5:LUCIFERASE reporter expression revealed that occasionally two pairs of LR founder cells are specified in close proximity even in wild-type and that one of them exists only transiently and disappears in an RLK7-dependent manner. We propose that the selection of LR founder cells by the peptide hormone-receptor cascade ensures proper LR spacing., (Copyright © 2018 Elsevier Inc. All rights reserved.)

Plants encode a unique group of papain-type cysteine endopeptidases (CysEP) characterized by a C-terminal KDEL endoplasmic reticulum retention signal (KDEL-CysEP) and an unusually broad substrate specificity. The three Arabidopsis KDEL-CysEPs (AtCEP1, AtCEP2, and AtCEP3) are differentially expressed in vegetative and generative tissues undergoing programmed cell death (PCD). While KDEL-CysEPs have been shown to be implicated in the collapse of tissues during PCD, roles of these peptidases in processes other than PCD are unknown. Using mCherry-AtCEP2 and EGFP-AtCEP1 reporter proteins in wild type versus atcep2 or atcep1 mutant plants, we explored the participation of AtCEP in young root development. Loss of AtCEP2, but not AtCEP1 resulted in shorter primary roots due to a decrease in cell length in the lateral root (LR) cap, and impairs extension of primary root epidermis cells such as trichoblasts in the elongation zone. AtCEP2 was localized to root cap corpses adherent to epidermal cells in the rapid elongation zone. AtCEP1 and AtCEP2 are expressed in root epidermis cells that are separated for LR emergence. Loss of AtCEP1 or AtCEP2 caused delayed emergence of LR primordia. KDEL-CysEPs might be involved in developmental tissue remodeling by supporting cell wall elongation and cell separation., Competing Interests: The authors have declared that no competing interests exist.