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Poplar is one of the fastest-growing temperate trees in the world and is widely used in ornamental horticulture for shade. The root is essential for tree growth and development and its utilization potential is huge. Calcium (Ca), as a signaling molecule, is involved in the regulation of plant root development. However, the detailed underlying regulatory mechanism is elusive. In this study, we analyzed the morphological and transcriptomic variations of 84K poplar (Populus alba x P. glandulosa) in response to different calcium concentrations and found that low Ca2+ (1 mmol • L-1) promoted lateral root development, while deficiency (0.1 mmol • L-1 Ca2+) inhibited lateral root development. Co-expression analysis showed that Ca2+ channel glutamate receptors (GLRs) were present in various modules with significance for root development. Two GLR paralogous genes, PagGLR3.3a and PagGLR3.3b, were mainly expressed in roots and up-regulated under Ca2+ deficiency. The CRISPR/Cas9-mediated signal gene (crispr-PagGLR3.3a, PagGLR3.3b) and double gene (crispr-PagGLR3.3ab) mutants presented more and longer lateral roots. Anatomical analysis showed that crispr-PagGLR3.3ab plants had more xylem cells and promoted the development of secondary vascular tissues. Further transcriptomic analysis suggested that knockout of PagGLR3.3a and PagGLR3.3b led to the up-regulation of several genes related to protein phosphorylation, auxin efflux, lignin and hemicellulose biosynthesis as well as transcriptional regulation, which might contribute to lateral root growth. This study not only provides novel insight into how the Ca2+ channels mediated root growth and development in trees, but also provides a directive breeding of new poplar species for biofuel and bioenergy production. [ABSTRACT FROM AUTHOR]

Summary Root morphogenesis is crucial for water and nutrient acquisition, but many aspects of root morphogenesis in crops are not well‐understood. Here, we cloned and functionally characterized a key gene for root morphogenesis in rice (Oryza sativa) based on mutant analysis. The stop root morphogenesis 1 (srm1) mutant lacks crown roots (CRs) and lateral roots (LRs) and carries a point mutation in the t‐SNARE coding gene SYNTAXIN OF PLANTS 132 (OsSYP132), leading to a premature stop codon and ablating the post‐transmembrane (PTM) region of OsSYP132. We identified the functional SNARE complex OsSYP132–OsNPSN13–OsSYP71–OsVAMP721/722 and determined that the integrity of the PTM region of OsSYP132 is essential for OsSYP132‐based SNARE complex‐mediated fusion of OsVAMP721/722 vesicles with the plasma membrane. The loss of this region in srm1 disrupts the intercellular trafficking and plasma membrane localization of OsPIN1b, preventing proper auxin distribution in the primordia of CRs and LRs and inhibiting their outgrowth. [ABSTRACT FROM AUTHOR]

Root-knot nematodes induce the formation of feeding sites within the host roots and the relocation of auxin into galls results in abnormal lateral root growth. Here, we analyzed the changes in cucumber root architecture under Meloidogyne incognita stress and the distribution of auxin in these morphological and molecular root changes. The number of root tips significantly decreased, and regression analysis showed a positive relationship between the size of root galls and the numbers of nematodes in galls compared with the lateral roots on galls, emphasizing the effect of nematode parasitism on root development. Data generated via a promoter-reporter system using the transgenic hairy root system first characterized the auxin distribution during nematode parasitism in cucumber. Using DR5:GUS staining of root galls, we further detected the expression of CsPIN1 and CsAUX1, which regulate polar auxin transport. The results showed that both CsPIN1 and CsAUX1 were induced in galls, and the relative expression of the two genes significantly increased at 21 DAI. The TIBA treatment, which can disrupt polar auxin transport inhibited the numbers of cucumber root tips and total length following increasing concentration gradients. Moreover, the numbers of galls were significantly affected by TIBA treatment, which showed the vital role of auxin during nematode parasitism. Our findings suggest that the transportation of auxin plays an important role during gall formation and induces cucumber lateral root development within nematode feeding sites. [ABSTRACT FROM AUTHOR]

Mediator, a multiprotein complex, is an important component of the transcription machinery. In plants, the latest studies have established that it functions as a signal processor that conveys transcriptional signals from transcription factors to RNA polymerase II. Mediator has been found to be involved in different developmental and stress-adaptation conditions, ranging from embryo, root, and shoot development to flowering and senescence, and also in responses to different biotic and abiotic stresses. In the last decade, significant progress has been made in understanding the role of Mediator subunits in root development. They have been shown to transcriptionally regulate development of almost all the components of the root system architecture—primary root, lateral roots, and root hairs. They also have a role in nutrient acquisition by the root. In this review, we discuss all the known functions of Mediator subunits during root development. We also highlight the role of Mediator as a nodal point for processing different hormone signals that regulate root morphogenesis and growth. [ABSTRACT FROM AUTHOR]

Rapeseed (Brassica napus) is an important oilseed crop worldwide. Plant vascular tissues are responsible for long-distance transport of water and nutrients and for providing mechanical support. The lateral roots absorb water and nutrients. The genetic basis of vascular tissue and lateral root development in rapeseed remains unknown. This study characterized an ethyl methanesulfonate-mutagenized rapeseed mutant, T16 , which showed dwarf stature, reduced lateral roots, and leaf wilting. SEM observations showed that the internode cells were shortened. Observations of tissue sections revealed defects in vascular bundle development in the stems and petioles. Genetic analysis revealed that the phenotypes of T16 were controlled by a single semi-dominant nuclear gene. Map-based cloning and genetic complementarity identified BnaA03.IAA13 as the functional gene; a G-to-A mutation in the second exon changed glycine at position 79 to glutamic acid, disrupting the conserved degron motif VGWPP. Transcriptome analysis in roots and stems showed that auxin and cytokinin signaling pathways were disordered in T16. Evolutionary analysis showed that AUXIN/INDOLE-3-ACETIC ACID is conserved during plant evolution. The heterozygote of T16 showed significantly reduced plant height while maintaining other agronomic traits. Our findings provide novel insights into the regulatory mechanisms of vascular tissue and lateral root development, and offer a new germplasm resource for rapeseed breeding. [ABSTRACT FROM AUTHOR]

Nitrate is a significant source of nitrogen in soils and also serves as a critical signal for root development. Previous studies have demonstrated that the local nitrate supply promotes lateral root elongation primarily through local nitrate signals, rather than nutritional effects. In this study, we report that Calcineurin B-like protein 1 (OsCBL1) positively regulates local nitrate signaling, thereby triggering lateral root colonization, as revealed by a comparative analysis of the phenotype and whole transcriptome of the knockdown mutant (OsCBL1-KD) and the wildtype (WT). In the split-root system, the knockdown of OsCBL1 was found to inhibit local nitrate-induced lateral root growth. Transcriptome analyses identified 398 differentially expressed genes (DEGs) that were under the control of OsCBL1 and associated with the phenotype of nitrate-induced lateral root colonization. Further analysis revealed that the nitrate transporter/sensor gene OsNRT1.1B was up-regulated under Sp-NaNO3 conditions compared to Sp-NaCl in WT but not in OsCBL1-KD plants. Pathway mapping of DEGs (i.e., genes exhibiting a significant change in expression in the Sp-NaNO3 condition compared to the Sp-NaCl condition) revealed a preferential upregulation of genes involved in lignin biosynthesis and a downregulation of genes involved in auxin and salicylic acid signaling. This suggests that OsCBL1 might function as a transmitter within the auxin, salicylic acid signaling, lignin biosynthesis, and nitrate sensor (OsNRT1.1B)-mediated pathways in response to local nitrate signaling. We also identified a transcriptional regulatory network downstream of OsCBL1 in nitrate-rich patches that is centered on several core transcription factors. Our study provides new insights into how plants adapt to an inhomogeneous distribution of nitrogen in the soil. [ABSTRACT FROM AUTHOR]

Summary Plant root system is significantly influenced by high soil levels of ammonium nitrogen, leading to reduced root elongation and enhanced lateral root branching. In Arabidopsis, these processes have been reported to be mediated by phytohormones and their downstream signaling pathways, while the controlling mechanisms remain elusive in crops. Through a transcriptome analysis of roots subjected to high/low ammonium treatments, we identified a cytokinin oxidase/dehydrogenase encoding gene, CKX3, whose expression is induced by high ammonium. Knocking out CKX3 and its homologue CKX8 results in shorter seminal roots, fewer lateral roots, and reduced sensitivity to high ammonium. Endogenous cytokinin levels are elevated by high ammonium or in ckx3 mutants. Cytokinin application results in shorter seminal roots and fewer lateral roots in wild‐type, mimicking the root responses of ckx3 mutants to high ammonium. Furthermore, CKX3 is transcriptionally activated by type‐B RR25 and RR26, and ckx3 mutants have reduced auxin content and signaling in roots under low ammonium. This study identified RR25/26‐CKX3‐cytokinin as a signal module that mediates root responses to external ammonium by modulating of auxin signaling in the root meristem and lateral root primordium. This highlights the critical role of cytokinin metabolism in regulating rice root development in response to ammonium. [ABSTRACT FROM AUTHOR]

The crucial role of TIR1‐receptor‐mediated gene transcription regulation in auxin signaling has long been established. In recent years, the significant role of protein phosphorylation modifications in auxin signal transduction has gradually emerged. To further elucidate the significant role of protein phosphorylation modifications in auxin signaling, a phosphoproteomic analysis in conjunction with auxin treatment has identified an auxin activated Mitogen‐activated Protein Kinase Kinase Kinase (MAPKKK) VH1‐INTERACTING Kinase (VIK), which plays an important role in auxin‐induced lateral root (LR) development. In the vik mutant, auxin‐induced LR development is significantly attenuated. Further investigations show that VIK interacts separately with the positive regulator of LR development, LATERAL ORGAN BOUNDARIES‐DOMAIN18 (LBD18), and the negative regulator of LR emergence, Ethylene Responsive Factor 13 (ERF13). VIK directly phosphorylates and stabilizes the positive transcription factor LBD18 in LR formation. In the meantime, VIK directly phosphorylates the negative regulator ERF13 at Ser168 and Ser172 sites, causing its degradation and releasing the repression by ERF13 on LR emergence. In summary, VIK‐mediated auxin signaling regulates LR development by enhancing the protein stability of LBD18 and inducing the degradation of ERF13, respectively. [ABSTRACT FROM AUTHOR]

The lateral root (LR) is an essential component of the plant root system, performing important functions for nutrient and water uptake in plants and playing a pivotal role in cereal crop productivity. Nitrate (NO3−) is an essential nutrient for plants. In this study, wheat plants were grown in 1/2 strength Hoagland's solution containing 5 mM NO3− (check; CK), 0.1 mM NO3− (low NO3−; LN), or 0.1 mM NO3− plus 60 mg/L 2,3,5-triiodobenzoic acid (TIBA) (LNT). The results showed that LN increased the LR number significantly at 48 h after treatment compared with CK, while not increasing the root biomass, and LNT significantly decreased the LR number and root biomass. The transcriptomic analysis showed that LN induced the expression of genes related to root IAA synthesis and transport and cell wall remodeling, and it was suppressed in the LNT conditions. A physiological assay revealed that the LN conditions increased the activity of IAA biosynthesis-related enzymes, the concentrations of tryptophan and IAA, and the activity of cell wall remodeling enzymes in the roots, whereas the content of polysaccharides in the LRP cell wall was significantly decreased compared with the control. Fourier-transform infrared spectroscopy and atomic microscopy revealed that the content of cell wall polysaccharides decreased and the cell wall elasticity of LR primordia (LRP) increased under the LN conditions. The effects of LN on IAA synthesis and polar transport, cell wall remodeling, and LR development were abolished when TIBA was applied. Our findings indicate that NO3− starvation may improve auxin homeostasis and the biological properties of the LRP cell wall and thus promote LR initiation, while TIBA addition dampens the effects of LN on auxin signaling, gene expression, physiological processes, and the root architecture. [ABSTRACT FROM AUTHOR]

Main conclusion: TaZAT8-5B, a C2H2 zinc finger protein transcription factor, positively regulates drought tolerance in transgenic Arabidopsis. It promotes root growth under drought stress via the Aux/IAA-ARF module in the auxin signaling pathway. C2H2 zinc finger proteins (C2H2-ZFPs) represent the largest but relatively unexplored family of transcription factors in plants. This is particularly evident in wheat, where the functions of only a few C2H2-ZFP genes have been confirmed. In this study, we identified a novel C2H2-ZFP gene, TaZAT8-5B. This gene shows high expression in roots and flowers and is significantly induced by heat, drought, and salt stress. Under drought stress, overexpressing TaZAT8-5B in Arabidopsis resulted in increased proline content and superoxide dismutase (SOD) activity in leaves. It also led to reduced stomatal aperture and water loss, while inducing the expression of P5CS1, RD29A, and DREB1A. Consequently, it alleviated drought stress-induced malondialdehyde (MDA) accumulation and improved drought tolerance. Additionally, TaZAT8-5B promoted lateral root initiation under mannitol stress and enhanced both lateral and primary root growth under long-term drought stress. Moreover, TaZAT8-5B was induced by indole-3-acetic acid (IAA). Overexpressing TaZAT8-5B under drought stress significantly inhibited the expression of auxin signaling negative regulatory genes IAA12 and IAA14. Conversely, downstream genes (ARF7, LBD16, LBD18, and CDKA1) of IAA14 and IAA12 were upregulated in TaZAT8-5B overexpressing plants compared to wild-type (WT) plants. These findings suggest that TaZAT8-5B regulates root growth and development under drought stress via the Aux/IAA-ARF module in the auxin signaling pathway. In summary, this study elucidates the role of TaZAT8-5B in enhancing drought tolerance and its involvement in root growth and development through the auxin signaling pathway. These findings offer new insights into the functional analysis of homologous genes of TaZAT8-5B, particularly in Gramineae species. [ABSTRACT FROM AUTHOR]

Abstract Background Salt is an important factor that affects crop productivity. Plant hexokinases (HXKs) are key enzymes in the glycolytic pathway and sugar signaling transduction pathways of plants. In previous studies, we identified and confirmed the roles of GmHXK2 in salt tolerance. Results In this study, we analyzed the tissue-specific expression of GmHXK2 at different growth stages throughout the plant’s life cycle. The results showed that GmHXK2 was expressed significantly in all tissues at vegetative stages, including germination and seedling. However, no expression was detected in the pods, and there was little expression in flowers during the later mature period. Arabidopsis plants overexpressing the GmHXK2 (OE) had more lateral roots. The OE seedlings also produced higher levels of auxin and ascorbic acid (AsA). Additionally, the expression levels of genes PMM, YUC4/YUC6/YUC8, and PIN/LAX1,LAX3, which are involved respectively in the synthesis of AsA and auxin, as well as polar auxin transport, were upregulated in OE plants. This upregulation occurred specifically under exogenous glucose treatment. AtHKT1, AtSOS1, and AtNHX1 were up-regulated in OE plants under salt stress, suggesting that GmHXK2 may modulate salt tolerance by maintaining ion balance within the cells and alleviating damage caused by salt stress. Additionally, we further confirmed the interaction between GmHXK2 and the protein GmPMM through yeast two-hybridization and bimolecular fluorescence complementation assays, respectively. Conclusion The expression of GmHXK2 gene in plants is organ-specific and developmental stage specific. GmHXK2 not only regulates the synthesis of AsA and the synthesis and distribution of auxin, but also promotes root elongation and induces lateral root formation, potentially enhancing soil water absorption. This study reveals the crosstalk between sugar signaling and hormone signaling in plants, where GmHXK2 acts as a glucose sensor through its interaction with GmPMM, and sheds light on the molecular mechanism by which GmHXK2 gene is involved in salt tolerance in plants.

Nitrogen (N) is an essential mineral element for plants and is the main component of protein, nucleic acid, phospholipid, chlorophyll, hormones, vitamins and alkaloids. It is involved in all stages of plant growth and development. Low-N stress seriously hinders plant growth and reduces yield and quality. Plants have evolved a series of elaborate regulatory mechanisms for N uptake and assimilation to cope with different soil N states. After N is absorbed and utilized by plants, it plays important regulatory roles in phytohormones, microRNA (miRNA), root development and mycorrhizal symbiosis to cope with environmental stress. Here, we highlight the research progress on the regulation mechanisms for N absorption and assimilation. Then, we emphasize the regulation roles of N regarding hormone signals, miRNA, lateral root growth, drought resistance, anthocyanin synthesis and mycorrhizal symbiosis. A thorough understanding of N uptake, utilization and interaction with other biological processes is helpful to improve N use efficiency and to breed plants with "less-input-more-output". [ABSTRACT FROM AUTHOR]

Background: Salt is an important factor that affects crop productivity. Plant hexokinases (HXKs) are key enzymes in the glycolytic pathway and sugar signaling transduction pathways of plants. In previous studies, we identified and confirmed the roles of GmHXK2 in salt tolerance. Results: In this study, we analyzed the tissue-specific expression of GmHXK2 at different growth stages throughout the plant's life cycle. The results showed that GmHXK2 was expressed significantly in all tissues at vegetative stages, including germination and seedling. However, no expression was detected in the pods, and there was little expression in flowers during the later mature period. Arabidopsis plants overexpressing the GmHXK2 (OE) had more lateral roots. The OE seedlings also produced higher levels of auxin and ascorbic acid (AsA). Additionally, the expression levels of genes PMM, YUC4/YUC6/YUC8, and PIN/LAX1,LAX3, which are involved respectively in the synthesis of AsA and auxin, as well as polar auxin transport, were upregulated in OE plants. This upregulation occurred specifically under exogenous glucose treatment. AtHKT1, AtSOS1, and AtNHX1 were up-regulated in OE plants under salt stress, suggesting that GmHXK2 may modulate salt tolerance by maintaining ion balance within the cells and alleviating damage caused by salt stress. Additionally, we further confirmed the interaction between GmHXK2 and the protein GmPMM through yeast two-hybridization and bimolecular fluorescence complementation assays, respectively. Conclusion: The expression of GmHXK2 gene in plants is organ-specific and developmental stage specific. GmHXK2 not only regulates the synthesis of AsA and the synthesis and distribution of auxin, but also promotes root elongation and induces lateral root formation, potentially enhancing soil water absorption. This study reveals the crosstalk between sugar signaling and hormone signaling in plants, where GmHXK2 acts as a glucose sensor through its interaction with GmPMM, and sheds light on the molecular mechanism by which GmHXK2 gene is involved in salt tolerance in plants. [ABSTRACT FROM AUTHOR]

Lateral roots (LRs) receive signals from the inter-root environment and absorb water and nutrients from the soil. Auxin regulates LR formation, but the mechanism in tomato remains largely unknown. In this study, 'Ailsa Craig' tomato LRs appeared on the third day and were unevenly distributed in primary roots. According to the location of LR occurrence, roots were divided into three equal parts: the shootward part of the root (RB), the middle part of the root (RM), and the tip part of the root (RT). Transverse sections of roots from days 1 to 6 revealed that the number of RB cells and the root diameter were significantly increased compared with RM and RT. Using roots from days 1 to 3, we carried out transcriptome sequencing analysis. Identified genes were classified into 16 co-expression clusters based on K-means, and genes in four associated clusters were highly expressed in RB. These four clusters (3, 5, 8, and 16) were enriched in cellulose metabolism, microtubule, and peptide metabolism pathways, all closely related to LR development. The four clusters contain numerous transcription factors linked to LR development including transcription factors of LATERAL ORGAN BOUNDRIES (LOB) and MADS-box families. Additionally, auxin-related genes GATA23, ARF7, LBD16, EXP, IAA4, IAA7, PIN1, PIN2, YUC3, and YUC4 were highly expressed in RB tissue. Free IAA content in 3 d RB was notably higher, reaching 3.3–5.5 ng/g, relative to RB in 1 d and 2 d. The LR number was promoted by 0.1 μM of exogenous IAA and inhibited by exogenous NPA. We analyzed the root cell state and auxin signaling module during LR formation. At a certain stage of pericycle cell development, LR initiation is regulated by auxin signaling modules IAA14-ARF7/ARF19-LBD16-CDKA1 and IAA14-ARF7/ARF19-MUS/MUL-XTR6/EXP. Furthermore, as a key regulatory factor, auxin regulates the process of LR initiation and LR primordia (LRP) through different auxin signaling pathway modules. [ABSTRACT FROM AUTHOR]

Auxin dictates root architecture via the Auxin Response Factor (ARF) family of transcription factors, which control lateral root (LR) formation. In Arabidopsis, ARF7 regulates the specification of prebranch sites (PBS) generating LRs through gene expression oscillations and plays a pivotal role during LR initiation. Despite the importance of ARF7 in this process, there is a surprising lack of knowledge about how ARF7 turnover is regulated and how this impacts root architecture. Here, we show that ARF7 accumulates in autophagy mutants and is degraded through NBR1-dependent selective autophagy. We demonstrate that the previously reported rhythmic changes to ARF7 abundance in roots are modulated via autophagy and might occur in other tissues. In addition, we show that the level of co-localization between ARF7 and autophagy markers oscillates and can be modulated by auxin to trigger ARF7 turnover. Furthermore, we observe that autophagy impairment prevents ARF7 oscillation and reduces both PBS establishment and LR formation. In conclusion, we report a novel role for autophagy during development, namely by enacting auxin-induced selective degradation of ARF7 to optimize periodic root branching. Synopsis: Auxin triggers NBR1-mediated selective autophagic degradation of ARF7 to modulate periodic root branching in Arabidopsis. ARF7 is selectively degraded via NBR1-mediated autophagy Auxin regulates ARF7 co-localization with autophagosomes and subsequent turnover Defective autophagy interferes with the root clock function and lateral root formation. Auxin triggers NBR1-mediated selective autophagic degradation of ARF7 to modulate periodic root branching in Arabidopsis. [ABSTRACT FROM AUTHOR]

Rhizobia induces nitrogen-fixing nodules in legumes used in agricultural production, providing a direct source of combined nitrogen to leguminous crops. Small peptides, such as CLAVATA3/EMBRYO SURROUNDING REGION peptides (CLE), are known to regulate the formation and development of nitrogen-fixing nodules in legumes. Root meristem growth factor (RGF) peptides from Medicago truncatula not only regulate root development but also modulate nodulation symbiosis with Sinorhizobium meliloti. However, the impact of RGF peptides from one leguminous species on the others remains unclear. In this study, we investigate the effects of the RGF family peptide MtRGF6p from M. truncatula on nodulation symbiosis and root development in Lotus japonicus. The MtRGF6 gene is predominantly expressed in the root nodules of M. truncatula and shows low identity with RGF homologous genes from L. japonicus. The gene promoter is active in the primordia of root nodules and lateral roots, as well as in young nodules and roots, and the meristem, infection, and nitrogen-fixing regions of the mature nodule. Chemically synthesized MtRGF6p promoted primary root growth in M. truncatula but suppressed the growth of L. japonicus primary roots. The peptide negatively affected the initiation of nodule primordia, the formation of infection threads, and nodulation in both legumes, with a low dosage showing effects on L. japonicus compared to M. truncatula. These results suggest that the MtRGF6 peptide from M. truncatula may serve as an inter-species signal affecting the root organ development of L. japonicus. [ABSTRACT FROM AUTHOR]

The phytohormone auxin triggers root growth inhibition within seconds via a non-transcriptional pathway. Among members of the TIR1/AFB auxin receptor family, AFB1 has a primary role in this rapid response. However, the unique features that confer this specific function have not been identified. Here we show that the N-terminal region of AFB1, including the F-box domain and residues that contribute to auxin binding, is essential and sufficient for its specific role in the rapid response. Substitution of the N-terminal region of AFB1 with that of TIR1 disrupts its distinct cytoplasm-enriched localization and activity in rapid root growth inhibition by auxin. Importantly, the N-terminal region of AFB1 is indispensable for auxin-triggered calcium influx, which is a prerequisite for rapid root growth inhibition. Furthermore, AFB1 negatively regulates lateral root formation and transcription of auxin-induced genes, suggesting that it plays an inhibitory role in canonical auxin signaling. These results suggest that AFB1 may buffer the transcriptional auxin response, whereas it regulates rapid changes in cell growth that contribute to root gravitropism.

Abstract The crucial role of TIR1‐receptor‐mediated gene transcription regulation in auxin signaling has long been established. In recent years, the significant role of protein phosphorylation modifications in auxin signal transduction has gradually emerged. To further elucidate the significant role of protein phosphorylation modifications in auxin signaling, a phosphoproteomic analysis in conjunction with auxin treatment has identified an auxin activated Mitogen‐activated Protein Kinase Kinase Kinase (MAPKKK) VH1‐INTERACTING Kinase (VIK), which plays an important role in auxin‐induced lateral root (LR) development. In the vik mutant, auxin‐induced LR development is significantly attenuated. Further investigations show that VIK interacts separately with the positive regulator of LR development, LATERAL ORGAN BOUNDARIES‐DOMAIN18 (LBD18), and the negative regulator of LR emergence, Ethylene Responsive Factor 13 (ERF13). VIK directly phosphorylates and stabilizes the positive transcription factor LBD18 in LR formation. In the meantime, VIK directly phosphorylates the negative regulator ERF13 at Ser168 and Ser172 sites, causing its degradation and releasing the repression by ERF13 on LR emergence. In summary, VIK‐mediated auxin signaling regulates LR development by enhancing the protein stability of LBD18 and inducing the degradation of ERF13, respectively.

Although strongly influenced by environmental conditions, lateral root (LR) positioning along the primary root appears to follow obediently an internal spacing mechanism dictated by auxin oscillations that prepattern the primary root, referred to as the root clock. Surprisingly, none of the hitherto characterized PIN- and ABCB-type auxin transporters seem to be involved in this LR prepatterning mechanism. Here, we characterize ABCB15, 16, 17, 18, and 22 (ABCB15-22) as novel auxin-transporting ABCBs. Knock-down and genome editing of this genetically linked group of ABCBs caused strongly reduced LR densities. These phenotypes were correlated with reduced amplitude, but not reduced frequency of the root clock oscillation. High-resolution auxin transport assays and tissue-specific silencing revealed contributions of ABCB15-22 to shootward auxin transport in the lateral root cap (LRC) and epidermis, thereby explaining the reduced auxin oscillation. Jointly, these data support a model in which LRC-derived auxin contributes to the root clock amplitude.

SUMMARY: The conservation of GOLVEN (GLV)/ROOT MERISTEM GROWTH FACTOR (RGF) peptide encoding genes across plant genomes capable of forming roots or root‐like structures underscores their potential significance in the terrestrial adaptation of plants. This study investigates the function and role of GOLVEN peptide‐coding genes in Medicago truncatula. Five out of fifteen GLV/RGF genes were notably upregulated during nodule organogenesis and were differentially responsive to nitrogen deficiency and auxin treatment. Specifically, the expression of MtGLV9 and MtGLV10 at nodule initiation sites was contingent upon the NODULE INCEPTION transcription factor. Overexpression of these five nodule‐induced GLV genes in hairy roots of M. truncatula and application of their synthetic peptide analogues led to a decrease in nodule count by 25–50%. Uniquely, the GOLVEN10 peptide altered the positioning of the first formed lateral root and nodule on the primary root axis, an observation we term 'noduletaxis'; this decreased the length of the lateral organ formation zone on roots. Histological section of roots treated with synthetic GOLVEN10 peptide revealed an increased cell number within the root cortical cell layers without a corresponding increase in cell length, leading to an elongation of the root likely introducing a spatiotemporal delay in organ formation. At the transcription level, the GOLVEN10 peptide suppressed expression of microtubule‐related genes and exerted its effects by changing expression of a large subset of Auxin responsive genes. These findings advance our understanding of the molecular mechanisms by which GOLVEN peptides modulate root morphology, nodule ontogeny, and interactions with key transcriptional pathways. Significance Statement: Nodule positioning is an understudied trait, yet it determines the length of the root that can support nodule formation and consequently the total number of functional nodules formed. We identify genetic factors called GOLVEN peptides that alter nodule and lateral root positioning on the primary root along with several other traits including nodule organ initiation and root architecture. [ABSTRACT FROM AUTHOR]

MicroRNAs (miRNAs) are a class of endogenous, non-coding small-molecule RNAs that usually regulate the expression of target genes at the post-transcriptional level. miR156 is one of a class of evolutionarily highly conserved miRNA families. SQUAMOSA PROMOTER BINDING PROTEIN-LIKE (SPL) transcription factor is one of the target genes that is regulated by miR156. SPL transcription factors are involved in regulating plant growth and development, hormone response, stress response, and photosynthesis. In the present study, transgenic potato plants with overexpressed miR156 were obtained via the Agrobacterium-mediated transformation method. The results showed that the expression levels of the target gene, StSPL9, were all downregulated in the transgenic plants with overexpressed Stu-miR156. Compared with those of the control plants, the plant height and root length of the transgenic plants were significantly decreased, while the number of lateral roots was significantly increased. These results revealed that the miR156/SPLs module was involved in regulating potato plant height and root growth. [ABSTRACT FROM AUTHOR]

Background: This study of the human brachial plexus quantifies and compares the total number of fibers, fiber diameter, and fiber density within the lateral and medial roots of the median nerve in twenty-seven adult human cadavers. The lateral and medial cords give rise to the lateral and medial roots of median nerve, which join to form the median nerve. In this study, the fiber ratios of small-diameter Group III and IV sensory fibers as well as large-diameter Group I and II sensory fibers comprising the lateral and medial roots of the median nerve of the brachial plexus were investigated. Methods: Peripheral nervous tissue sections of twenty-seven lateral and twenty-seven medial roots were surgically removed from the axilla region of formalin-fixed cadavers in 2021. Excised tissue sections were fractured using liquid nitrogen-based freeze-cracking, and subsequently viewed using scanning electron microscopy. ImageJ 1.x, Microsoft Excel, and Statistica software were used to analyze fiber count, fiber diameter, fiber density, and fascicular bundle count. Results: A higher ratio of large-diameter Group I and II sensory fibers comprise the lateral root, and a higher ratio of small-diameter Group III and IV fibers comprise the medial root. Conclusions: Understanding the fiber composition of the lateral and medial roots of the median nerve can aid in interpreting why avulsive injury of the upper or lower roots of the brachial plexus can lead to different pain experiences post-injury.

Root-knot nematodes induce the formation of feeding sites within the host roots and the relocation of auxin into galls results in abnormal lateral root growth. Here, we analyzed the changes in cucumber root architecture under Meloidogyne incognita stress and the distribution of auxin in these morphological and molecular root changes. The number of root tips significantly decreased, and regression analysis showed a positive relationship between the size of root galls and the numbers of nematodes in galls compared with the lateral roots on galls, emphasizing the effect of nematode parasitism on root development. Data generated via a promoter-reporter system using the transgenic hairy root system first characterized the auxin distribution during nematode parasitism in cucumber. Using DR5:GUS staining of root galls, we further detected the expression of CsPIN1 and CsAUX1, which regulate polar auxin transport. The results showed that both CsPIN1 and CsAUX1 were induced in galls, and the relative expression of the two genes significantly increased at 21 DAI. The TIBA treatment, which can disrupt polar auxin transport inhibited the numbers of cucumber root tips and total length following increasing concentration gradients. Moreover, the numbers of galls were significantly affected by TIBA treatment, which showed the vital role of auxin during nematode parasitism. Our findings suggest that the transportation of auxin plays an important role during gall formation and induces cucumber lateral root development within nematode feeding sites.

The lateral root (LR) is an essential component of the plant root system, performing important functions for nutrient and water uptake in plants and playing a pivotal role in cereal crop productivity. Nitrate (NO3−) is an essential nutrient for plants. In this study, wheat plants were grown in 1/2 strength Hoagland’s solution containing 5 mM NO3− (check; CK), 0.1 mM NO3− (low NO3−; LN), or 0.1 mM NO3− plus 60 mg/L 2,3,5-triiodobenzoic acid (TIBA) (LNT). The results showed that LN increased the LR number significantly at 48 h after treatment compared with CK, while not increasing the root biomass, and LNT significantly decreased the LR number and root biomass. The transcriptomic analysis showed that LN induced the expression of genes related to root IAA synthesis and transport and cell wall remodeling, and it was suppressed in the LNT conditions. A physiological assay revealed that the LN conditions increased the activity of IAA biosynthesis-related enzymes, the concentrations of tryptophan and IAA, and the activity of cell wall remodeling enzymes in the roots, whereas the content of polysaccharides in the LRP cell wall was significantly decreased compared with the control. Fourier-transform infrared spectroscopy and atomic microscopy revealed that the content of cell wall polysaccharides decreased and the cell wall elasticity of LR primordia (LRP) increased under the LN conditions. The effects of LN on IAA synthesis and polar transport, cell wall remodeling, and LR development were abolished when TIBA was applied. Our findings indicate that NO3− starvation may improve auxin homeostasis and the biological properties of the LRP cell wall and thus promote LR initiation, while TIBA addition dampens the effects of LN on auxin signaling, gene expression, physiological processes, and the root architecture.

The architecture of the root system is fundamental to plant productivity. The rate of root growth, the density of lateral roots, and the spatial structure of lateral and adventitious roots determine the developmental plasticity of the root system in response to changes in environmental conditions. One of the genes involved in the regulation of the slope angle of lateral roots is DEEPER ROOTING 1 (DRO1). Its orthologs and paralogs have been identified in rice, Arabidopsis, and several other species. However, nothing is known about the formation of the slope angle of lateral roots in species with the initiation of lateral root primordia within the parental root meristem. To address this knowledge gap, we identified orthologs and paralogs of the DRO1 gene in cucumber (Cucumis sativus) using a phylogenetic analysis of IGT protein family members. Differences in the transcriptional response of CsDRO1, CsDRO1-LIKE1 (CsDRO1L1), and CsDRO1-LIKE2 (CsDRO1L2) to exogenous auxin were analyzed. The results showed that only CsDRO1L1 is auxin-responsive. An analysis of promoter–reporter fusions demonstrated that the CsDRO1, CsDRO1L1, and CsDRO1L2 genes were expressed in the meristem in cell files of the central cylinder, endodermis, and cortex; the three genes displayed different expression patterns in cucumber roots with only partial overlap. A knockout of individual CsDRO1, CsDRO1L1, and CsDRO1L2 genes was performed via CRISPR/Cas9 gene editing. Our study suggests that the knockout of individual genes does not affect the slope angle formation during lateral root primordia development in the cucumber parental root. [ABSTRACT FROM AUTHOR]

Plant roots fulfil crucial tasks during a plant's life. As roots encounter very diverse conditions while exploring the soil for resources, their growth and development must be responsive to changes in the rhizosphere, resulting in root architectures that are tailor-made for all prevailing circumstances. Using multi-disciplinary approaches, we are gaining more intricate insights into the regulatory mechanisms directing root system architecture. This Special Issue provides insights into our advancement of knowledge on different aspects of root development and identifies opportunities for future research. [ABSTRACT FROM AUTHOR]

Plant submergence is a major abiotic stress that impairs plant performance. Under water, reduced gas diffusion exposes submerged plant cells to an environment that is enriched in gaseous ethylene and is limited in oxygen (O2) availability (hypoxia). The capacity for plant roots to avoid and/or sustain critical hypoxia damage is essential for plants to survive waterlogging. Plants use spatiotemporal ethylene and O2 dynamics as instrumental flooding signals to modulate potential adaptive root growth and hypoxia stress acclimation responses. However, how non-adapted plant species modulate root growth behaviour during actual waterlogged conditions to overcome flooding stress has hardly been investigated. Here we discuss how changes in the root growth rate, lateral root formation, density, and growth angle of non-flood adapted plant species (mainly Arabidopsis) could contribute to avoiding and enduring critical hypoxic conditions. In addition, we discuss current molecular understanding of how ethylene and hypoxia signalling control these adaptive root growth responses. We propose that future research would benefit from less artificial experimental designs to better understand how plant roots respond to and survive waterlogging. This acquired knowledge would be instrumental to guide targeted breeding of flood-tolerant crops with more resilient root systems. [ABSTRACT FROM AUTHOR]

Lateral roots are a major component of root system architecture, and lateral root count (LRC) positively contributes to yield under drought in chickpea. To understand the genetic regulation of LRC, a biparental mapping population derived from two chickpea accessions having contrasting LRCs was genotyped by sequencing, and phenotyped to map four major quantitative trait loci (QTLs) contributing to 13–32% of the LRC trait variation. A single- nucleotide polymorphism tightly linked to the locus contributing to highest trait variation was located on the coding region of a gene (CaWIP2), orthologous to NO TRANSMITTING TRACT/WIP domain protein 2 (NTT/WIP2) gene of Arabidopsis thaliana. A polymorphic simple sequence repeat (SSR) in the CaWIP2 promoter showed differentiation between low versus high LRC parents and mapping individuals, suggesting its utility for marker-assisted selection. CaWIP2 promoter showed strong expression in chickpea apical root meristem and lateral root primordia. Expression of CaWIP2 under its native promoter in the Arabidopsis wip2wip4wip5 mutant rescued its rootless phenotype to produce more lateral roots than the wild-type plants, and led to formation of amyloplasts in the columella. CaWIP2 expression also induced the expression of genes that regulate lateral root emergence. Our study identified a gene-based marker for LRC which will be useful for developing drought-tolerant, high-yielding chickpea varieties. [ABSTRACT FROM AUTHOR]

Summary: Lateral root (LR) positioning and development rely on the dynamic interplay between auxin production, transport but also inactivation. Nonetheless, how the latter affects LR organogenesis remains largely uninvestigated.Here, we systematically analyze the impact of the major auxin inactivation pathway defined by GRETCHEN HAGEN3‐type (GH3) auxin conjugating enzymes and DIOXYGENASE FOR AUXIN OXIDATION1 (DAO1) in all stages of LR development using reporters, genetics and inhibitors in Arabidopsis thaliana.Our data demonstrate that the gh3.1/2/3/4/5/6 hextuple (gh3hex) mutants display a higher LR density due to increased LR initiation and faster LR developmental progression, acting epistatically over dao1‐1. Grafting and local inhibitor applications reveal that root and shoot GH3 activities control LR formation. The faster LR development in gh3hex is associated with GH3 expression domains in and around developing LRs. The increase in LR initiation is associated with accelerated auxin response oscillations coinciding with increases in apical meristem size and LR cap cell death rates.Our research reveals how GH3‐mediated auxin inactivation attenuates LR development. Local GH3 expression in LR primordia attenuates development and emergence, whereas GH3 effects on pre‐initiation stages are indirect, by modulating meristem activities that in turn coordinate root growth with LR spacing. [ABSTRACT FROM AUTHOR]

Given the consistent release of zinc oxide (ZnO) nanoparticles into the environment, it is urgent to study their impact on plants in depth. In this study, grains of rice were treated with two different concentrations of ZnO nanoparticles (NP-ZnO, 10 and 100 mg/L), and their bulk counterpart (B-ZnO) were used to evaluate whether ZnO action could depend on particle size. To test this hypothesis, root growth and development assessment, oxidative stress parameters, indole-3-acetic acid (IAA) content and molecules/enzymes involved in IAA metabolism were analyzed. In situ localization of Zn in control and treated roots was also performed. Though Zn was visible inside root cells only following nanoparticle treatment, both materials (NP-ZnO and B-ZnO) were able to affect seedling growth and root morphology, with alteration in the concentration/pattern of localization of oxidative stress markers and with a different action depending on particle size. In addition, only ZnO supplied as bulk material induced a significant increase in both IAA concentration and lateral root density, supporting our hypothesis that bulk particles might enhance lateral root development through the rise of IAA concentration. Apparently, IAA concentration was influenced more by the activity of the catabolic peroxidases than by the protective action of phenols.

Two sorghum varieties, Shanqui Red (SQR) and SRN39, have distinct levels of susceptibility to the parasitic weed Striga hermonthica, which have been attributed to different strigolactone composition within their root exudates. Root exudates of the Striga-susceptible variety Shanqui Red (SQR) contain primarily 5-deoxystrigol, which has a high efficiency for inducing Striga germination. SRN39 roots primarily exude orobanchol, leading to reduced Striga germination and making this variety resistant to Striga. The structural diversity in exuded strigolactones is determined by a polymorphism in the LOW GERMINATION STIMULANT 1 (LGS1) locus. Yet, the genetic diversity between SQR and SRN39 is broad and has not been addressed in terms of growth and development. Here, we demonstrate additional differences between SQR and SRN39 by phenotypic and molecular characterization. A suite of genes related to metabolism was differentially expressed between SQR and SRN39. Increased levels of gibberellin precursors in SRN39 were accompanied by slower growth rate and developmental delay and we observed an overall increased SRN39 biomass. The slow-down in growth and differences in transcriptome profiles of SRN39 were strongly associated with plant age. Additionally, enhanced lateral root growth was observed in SRN39 and three additional genotypes exuding primarily orobanchol. In summary, we demonstrate that the differences between SQR and SRN39 reach further than the changes in strigolactone profile in the root exudate and translate into alterations in growth and development.

Nitrogen (N) is an essential mineral element for plants and is the main component of protein, nucleic acid, phospholipid, chlorophyll, hormones, vitamins and alkaloids. It is involved in all stages of plant growth and development. Low-N stress seriously hinders plant growth and reduces yield and quality. Plants have evolved a series of elaborate regulatory mechanisms for N uptake and assimilation to cope with different soil N states. After N is absorbed and utilized by plants, it plays important regulatory roles in phytohormones, microRNA (miRNA), root development and mycorrhizal symbiosis to cope with environmental stress. Here, we highlight the research progress on the regulation mechanisms for N absorption and assimilation. Then, we emphasize the regulation roles of N regarding hormone signals, miRNA, lateral root growth, drought resistance, anthocyanin synthesis and mycorrhizal symbiosis. A thorough understanding of N uptake, utilization and interaction with other biological processes is helpful to improve N use efficiency and to breed plants with “less-input-more-output”.

Lateral roots (LRs) receive signals from the inter-root environment and absorb water and nutrients from the soil. Auxin regulates LR formation, but the mechanism in tomato remains largely unknown. In this study, ‘Ailsa Craig‘ tomato LRs appeared on the third day and were unevenly distributed in primary roots. According to the location of LR occurrence, roots were divided into three equal parts: the shootward part of the root (RB), the middle part of the root (RM), and the tip part of the root (RT). Transverse sections of roots from days 1 to 6 revealed that the number of RB cells and the root diameter were significantly increased compared with RM and RT. Using roots from days 1 to 3, we carried out transcriptome sequencing analysis. Identified genes were classified into 16 co-expression clusters based on K-means, and genes in four associated clusters were highly expressed in RB. These four clusters (3, 5, 8, and 16) were enriched in cellulose metabolism, microtubule, and peptide metabolism pathways, all closely related to LR development. The four clusters contain numerous transcription factors linked to LR development including transcription factors of LATERAL ORGAN BOUNDRIES (LOB) and MADS-box families. Additionally, auxin-related genes GATA23, ARF7, LBD16, EXP, IAA4, IAA7, PIN1, PIN2, YUC3, and YUC4 were highly expressed in RB tissue. Free IAA content in 3 d RB was notably higher, reaching 3.3–5.5 ng/g, relative to RB in 1 d and 2 d. The LR number was promoted by 0.1 μM of exogenous IAA and inhibited by exogenous NPA. We analyzed the root cell state and auxin signaling module during LR formation. At a certain stage of pericycle cell development, LR initiation is regulated by auxin signaling modules IAA14-ARF7/ARF19-LBD16-CDKA1 and IAA14-ARF7/ARF19-MUS/MUL-XTR6/EXP. Furthermore, as a key regulatory factor, auxin regulates the process of LR initiation and LR primordia (LRP) through different auxin signaling pathway modules.

Lateral roots (LRs) are an important part of plant root systems. In dicots, for example, after plants adapted from aquatic to terrestrial environments, filamentous pseudorhizae evolved to allow nutrient absorption. A typical plant root system comprises a primary root, LRs, root hairs, and a root cap. Classical plant roots exhibit geotropism (the tendency to grow downward into the ground) and can synthesize plant hormones and other essential substances. Root vascular bundles and complex spatial structures enable plants to absorb water and nutrients to meet their nutrient quotas and grow. The primary root carries out most functions during early growth stages but is later overtaken by LRs, underscoring the importance of LR development water and mineral uptake and the soil fixation capacity of the root. LR development is modulated by endogenous plant hormones and external environmental factors, and its underlying mechanisms have been dissected in great detail in Arabidopsis, thanks to its simple root anatomy and the ease of obtaining mutants. This review comprehensively and systematically summarizes past research (largely in Arabidopsis) on LR basic structure, development stages, and molecular mechanisms regulated by different factors, as well as future prospects in LR research, to provide broad background knowledge for root researchers.

Nodules are formed as a result of symbiosis with nitrogen-fixing bacteria on the roots of legume plants. Symbiotic nodule development shares common developmental processes with lateral roots, including cell proliferation of the pericycle and cortical cell layers leading to primordium formation, and both these organs are assumed to be evolutionarily related. The progression of their development is realized with the help of phytohormones, in particular auxin and cytokinin. Due to common evolutionary origin, the hormonal pathway and response may have similar functionality in cell fate specification during lateral root and nodule development. It is known that auxin and cytokinin both positively regulate nodule primordia development. However during lateral root development, auxin is a key positive regulator, while cytokinin suppresses lateral root development. Therefore, a comparative review of auxin and cytokinin action in the development of these two organs to elucidate how their identity is established should be of special interest. This review discusses the involvement of auxin and cytokinin in the development of symbiotic nodules and lateral roots, with a special focus on the similarity and differences of their action during lateral root and nodule developmental processes. The spatio-temporal peculiarities of auxin and cytokinin biosynthesis, transport and response during lateral root and symbiotic nodule formation are reviewed, which should help to clarify how the identity of these two root lateral organs depends on auxin-cytokinin interplay. [ABSTRACT FROM AUTHOR]

Plant roots are sensitive to potassium (K+) deficiency signals. Therefore, regulating root growth by exogenous methods is a vital strategy to improve low K+ tolerance of sweetpotato. We studied the effects of exogenous indole-3-acetic acid (IAA) on growth, K+ absorption, and root characteristics in sweetpotato exposed to low K+ treatment (LK). LK significantly inhibited dry mass, K+ concentration and accumulation, as well as the root elongation (length) and branching (forks and crossings) in sweetpotato seedlings. However, exogenous IAA increased the length, ratio, and density of lateral roots and promoted absorption and accumulation of K+, which effectively alleviated the inhibitory effect of low K+. Exogenous IAA also increased the expression levels of auxin synthesis (IbYUC6 and IbTAR2) and transport (IbPIN1, IbPIN3, and IbPIN8) genes in leaves and roots, which promoted the increase of endogenous IAA content. Furthermore, exogenous IAA was more effective on low-K-tolerant variety (XS32) than low-K-sensitive variety (NZ1) under LK stress, depending on their different IAA synthesis and transport strategies. These results indicated that exogenous IAA enhanced root responsiveness of sweetpotato to low K+ stress by modulating auxin biosynthesis and transport, thereby improving the tolerance of sweetpotato to low K+ stress. [ABSTRACT FROM AUTHOR]

Lateral root (LR) formation is an important developmental event for the establishment of the root system in most vascular plants. In Arabidopsis thaliana , the fewer roots (fwr) mutation in the GNOM gene, encoding a guanine nucleotide exchange factor of ADP ribosylation factor that regulates vesicle trafficking, severely inhibits LR formation. Local accumulation of auxin response for LR initiation is severely affected in fwr. To better understand how local accumulation of auxin response for LR initiation is regulated, we identified a mutation, fewer roots suppressor1 (fsp1), that partially restores LR formation in fwr. The gene responsible for fsp1 was identified as SUPERROOT2 (SUR2), encoding CYP83B1 that positions at the metabolic branch point in the biosynthesis of auxin/indole-3-acetic acid (IAA) and indole glucosinolate. The fsp1 mutation increases both endogenous IAA levels and the number of the sites where auxin response locally accumulates prior to LR formation in fwr. SUR2 is expressed in the pericycle of the differentiation zone and in the apical meristem in roots. Time-lapse imaging of the auxin response revealed that local accumulation of auxin response is more stable in fsp1. These results suggest that SUR2/CYP83B1 affects LR founder cell formation at the xylem pole pericycle cells where auxin accumulates. Analysis of the genetic interaction between SUR2 and GNOM indicates the importance of stabilization of local auxin accumulation sites for LR initiation. [ABSTRACT FROM AUTHOR]

Roots are the main organ for water uptake and the earliest part of a plant's response to drought, making them of great importance to our understanding of the root system's response to drought. However, little is known about the underlying molecular mechanisms that control root responses to drought stress. Here, we identified and functionally characterized the AP2/ERF family transcription factor (TF) PtrABR1 and the upstream target gene zinc-finger protein TF PtrYY1, which respond to drought stress by promoting the growth and development of lateral roots in Populus trichocarpa. A root-specific induction of PtrABR1 under drought stress was explored. The overexpression of PtrABR1 (PtrABR1-OE) promoted root growth and development, thereby increasing tolerance to drought stress. In addition, PtrYY1 is directly bound to the promoter of PtrABR1 under drought stress, and the overexpression of PtrYY1 (PtrYY1-OE) promoted lateral root growth and development and increased tolerance to drought stress. An RNA-seq analysis of PtrABR1-OE with wild-type (WT) poplar identified PtrGH3.6 and PtrPP2C44, which share the same pattern of expression changes as PtrABR1. A qRT-PCR and cis-element analysis further suggested that PtrGH3.6 and PtrPP2C44 may act as potential downstream targets of PtrABR1 genes in the root response pathway to drought stress. In conclusion, these results reveal a novel drought regulatory pathway in which PtrABR1 regulates the network through the upstream target gene PtrYY1 and the potential downstream target genes PtrGH3.6 and PtrPP2C44, thereby promoting root growth and development and improving tolerance to drought stress. [ABSTRACT FROM AUTHOR]

Given the consistent release of zinc oxide (ZnO) nanoparticles into the environment, it is urgent to study their impact on plants in depth. In this study, grains of rice were treated with two different concentrations of ZnO nanoparticles (NP-ZnO, 10 and 100 mg/L), and their bulk counterpart (B-ZnO) were used to evaluate whether ZnO action could depend on particle size. To test this hypothesis, root growth and development assessment, oxidative stress parameters, indole-3-acetic acid (IAA) content and molecules/enzymes involved in IAA metabolism were analyzed. In situ localization of Zn in control and treated roots was also performed. Though Zn was visible inside root cells only following nanoparticle treatment, both materials (NP-ZnO and B-ZnO) were able to affect seedling growth and root morphology, with alteration in the concentration/pattern of localization of oxidative stress markers and with a different action depending on particle size. In addition, only ZnO supplied as bulk material induced a significant increase in both IAA concentration and lateral root density, supporting our hypothesis that bulk particles might enhance lateral root development through the rise of IAA concentration. Apparently, IAA concentration was influenced more by the activity of the catabolic peroxidases than by the protective action of phenols. [ABSTRACT FROM AUTHOR]

MicroRNAs (miRNAs) are a class of endogenous, non-coding small-molecule RNAs that usually regulate the expression of target genes at the post-transcriptional level. miR156 is one of a class of evolutionarily highly conserved miRNA families. SQUAMOSA PROMOTER BINDING PROTEIN-LIKE (SPL) transcription factor is one of the target genes that is regulated by miR156. SPL transcription factors are involved in regulating plant growth and development, hormone response, stress response, and photosynthesis. In the present study, transgenic potato plants with overexpressed miR156 were obtained via the Agrobacterium-mediated transformation method. The results showed that the expression levels of the target gene, StSPL9, were all downregulated in the transgenic plants with overexpressed Stu-miR156. Compared with those of the control plants, the plant height and root length of the transgenic plants were significantly decreased, while the number of lateral roots was significantly increased. These results revealed that the miR156/SPLs module was involved in regulating potato plant height and root growth.

In low pH condition of soils (pH

Nutrient starvation-induced autophagy is a conserved process in eukaryotes. Plants defective in autophagy show hypersensitivity to carbon and nitrogen limitation. However, the role of autophagy in plant phosphate (Pi) starvation response is relatively less explored. Among the core autophagy-related (ATG) genes, ATG8 encodes a ubiquitin-like protein involved in autophagosome formation and selective cargo recruitment. The Arabidopsis thaliana ATG8 genes, AtATG8f and AtATG8h, are notably induced in roots under low Pi. In this study, we show that such upregulation correlates with their promoter activities and can be suppressed in the phosphate response 1 (phr1) mutant. Yeast one-hybrid analysis failed to attest the binding of the AtPHR1 transcription factor to the promoter regions of AtATG8f and AtATG8h. Dual luciferase reporter assays in Arabidopsis mesophyll protoplasts also indicated that AtPHR1 could not transactivate the expression of both genes. Loss of AtATG8f and AtATG8h leads to decreased root microsomal-enriched ATG8 but increased ATG8 lipidation. Moreover, atg8f/atg8h mutants exhibit reduced autophagic flux estimated by the vacuolar degradation of ATG8 in the Pi-limited root but maintain normal cellular Pi homeostasis with reduced number of lateral roots. While the expression patterns of AtATG8f and AtATG8h overlap in the root stele, AtATG8f is more strongly expressed in the root apex and root hair and remarkably at sites where lateral root primordia develop. We hypothesize that Pi starvation-induction of AtATG8f and AtATG8h may not directly contribute to Pi recycling but rely on a second wave of transcriptional activation triggered by PHR1 that fine-tunes cell type-specific autophagic activity. [ABSTRACT FROM AUTHOR]

The phytohormone auxin triggers root growth inhibition within seconds via a non-transcriptional pathway. Among members of the TIR1/AFB auxin receptor family, AFB1 has a primary role in this rapid response. However, the unique features that confer this specific function have not been identified. Here we show that the N-terminal region of AFB1, including the F-box domain and residues that contribute to auxin binding, is essential and sufficient for its specific role in the rapid response. Substitution of the N-terminal region of AFB1 with that of TIR1 disrupts its distinct cytoplasm-enriched localization and activity in rapid root growth inhibition by auxin. Importantly, the N-terminal region of AFB1 is indispensable for auxin-triggered calcium influx, which is a prerequisite for rapid root growth inhibition. Furthermore, AFB1 negatively regulates lateral root formation and transcription of auxin-induced genes, suggesting that it plays an inhibitory role in canonical auxin signaling. These results suggest that AFB1 may buffer the transcriptional auxin response, whereas it regulates rapid changes in cell growth that contribute to root gravitropism. The phytohormone auxin triggers ultra-rapid responses in Arabidopsis roots, such as a cytoplasmic calcium spike, enabling rapid gravitropic response of roots. This study reveals that the auxin receptor AFB1 controls these early auxin responses from the cytoplasm of root cells and negatively regulates the emergence of lateral roots. [ABSTRACT FROM AUTHOR]

Summary: An environmentally responsive root system is crucial for plant growth and crop yield, especially in suboptimal soil conditions. This responsiveness enables the plant to exploit regions of high nutrient density while simultaneously minimizing abiotic stress. Despite the vital importance of root systems in regulating plant growth, significant gaps of knowledge exist in the mechanisms that regulate their architecture.Auxin defines both the frequency of lateral root (LR) initiation and the rate of LR outgrowth. Here, we describe a search for proteins that regulate root system architecture (RSA) by interacting directly with a key auxin transporter, PIN1. The native separation of Arabidopsis plasma membrane protein complexes identified several PIN1 co‐purifying proteins.Among them, AZG1 was subsequently confirmed as a PIN1 interactor. Here, we show that, in Arabidopsis, AZG1 is a cytokinin (CK) import protein that co‐localizes with and stabilizes PIN1, linking auxin and CK transport streams. AZG1 expression in LR primordia is sensitive to NaCl, and the frequency of LRs is AZG1‐dependent under salt stress.This report therefore identifies a potential point for auxin:cytokinin crosstalk, which shapes RSA in response to NaCl. [ABSTRACT FROM AUTHOR]

Nitric oxide (NO) is involved not only in the regulation of plant growth, development, and stress responses but also in the regulation of plant-microbe interactions. Here, we demonstrate that Piriformospora indica can induce tobacco nitrate reductase to produce a NO signal in roots which enhances nitrogen uptake capacity by inducing the expression of ammonium and nitrate transporter genes and the development of lateral root and root hair, thereby promoting tobacco growth. In addition, the NO signal induced by P. indica is significantly different from that induced by the pathogen Phytophthora nicotianae. Inoculation with P. indica did not produce H2O2 and maintained high expression of Phytoglobin 1 in roots, resulting in a significantly lower NO level than in the roots inoculated with P. nicotianae. These findings suggest that an appropriate NO level is the likely basis of plant-P. indica symbiosis, which promotes the growth of host plants.

Plant organogenesis requires matching the available metabolic resources to developmental programs. In Arabidopsis, the root system is determined by primary root‐derived lateral roots (LRs), and adventitious roots (ARs) formed from non‐root organs. Lateral root formation entails the auxin‐dependent activation of transcription factors ARF7, ARF19, and LBD16. Adventitious root formation relies on LBD16 activation by auxin and WOX11. The allocation of shoot‐derived sugar to the roots influences branching, but how its availability is sensed for LRs formation remains unknown. We combine metabolic profiling with cell‐specific interference to show that LRs switch to glycolysis and consume carbohydrates. The target‐of‐rapamycin (TOR) kinase is activated in the lateral root domain. Interfering with TOR kinase blocks LR initiation while promoting AR formation. The target‐of‐rapamycin inhibition marginally affects the auxin‐induced transcriptional response of the pericycle but attenuates the translation of ARF19, ARF7, and LBD16. TOR inhibition induces WOX11 transcription in these cells, yet no root branching occurs as TOR controls LBD16 translation. TOR is a central gatekeeper for root branching that integrates local auxin‐dependent pathways with systemic metabolic signals, modulating the translation of auxin‐induced genes. Synopsis: Shoot‐derived sugar promotes lateral root formation, but the mechanism through which it influences cell differentiation in this context is not well understood. Here, we learn that TOR kinase mediates lateral root formation by inducing the translation of lateral root‐promoting transcription factor transcripts in a sugar‐dependent manner.Lateral root formation is an important consumer of shoot‐derived carbon.Pericycle activation requires shoot‐derived carbohydrates and TOR.TOR promotes the translation of transcription factors ARF19, ARF7, and LBD16.TOR suppresses non‐canonical primary root branching by inhibiting WOX11 expression. [ABSTRACT FROM AUTHOR]

Auxin distribution is essential for determining root developmental patterns. The formation of lateral roots and constitutive aerenchyma, which is a gas space developed through cell death, is regulated by auxin in rice (Oryza sativa). However, it is unclear whether the involvement of auxin in constitutive aerenchyma formation is conserved in other species. In this study, we found that constitutive aerenchyma formation was regulated by auxin in the nodal roots of Zea nicaraguensis, a wild relative of maize (Zea mays ssp. mays) grown naturally on frequently flooded coastal plains. Subsequent gravistimulation (root rotation) experiments showed opposite patterns of aerenchyma and lateral root formation. Lateral root formation on the convex side of rotated roots is known to be stimulated by a transient increase in auxin level in the pericycle. We found that aerenchyma formation was accelerated in the cortex on the concave side of the rotated nodal roots of Z. nicaraguensis. A cortex-specific expression analysis of auxin-responsive genes suggested that the auxin level was higher on the concave side than on the convex side. These results suggest that asymmetric auxin distribution underlies the regulation of aerenchyma and lateral root formation in the nodal roots of Z. nicaraguensis. As aerenchyma reduces the respiratory cost of the roots, constitutive aerenchyma on the concave side of the nodal root may balance resource allocation, thereby contributing to the uptake of water and nutrients by newly formed lateral roots. Our study provides insights into auxin-dependent asymmetric root patterning such as that of gravistimulation and hydropatterning response. [ABSTRACT FROM AUTHOR]

Summary: A strong root system facilitates the absorption of water and nutrients from the soil, to improve the growth of crops. However, to date, there are still very few root development regulatory genes that can be used in crop breeding for agriculture.In this study, we cloned a negative regulator gene of root development, Robust Root System 1 (RRS1), which encodes an R2R3‐type MYB family transcription factor. RRS1 knockout plants showed enhanced root growth, including longer root length, longer lateral root length, and larger lateral root density. RRS1 represses root development by directly activating the expression of OsIAA3 which is involved in the auxin signaling pathway.A natural variation in the coding region of RRS1 changes the transcriptional activity of its protein. RRS1T allele, originating from wild rice, possibly increases root length by means of weakening regulation of OsIAA3.Knockout of RRS1 enhances drought resistance by promoting water absorption and improving water use efficiency. This study provides a new gene resource for improving root systems and cultivating drought‐resistant rice varieties with important values in agricultural applications. [ABSTRACT FROM AUTHOR]

SUMMARY: Reactive oxygen species (ROS) play a dual role in plant biology, acting as important signal transduction molecules and as toxic byproducts of aerobic metabolism that accumulate in cells upon exposure to different stressors and lead to cell death. In plants, root architecture is regulated by the distribution and intercellular flow of the phytohormone auxin. In this study, we identified ROS as an important modulator of auxin distribution and response in the root. ROS production is necessary for root growth, proper tissue patterning, cell growth, and lateral root (LR) induction. Alterations in ROS balance led to altered auxin distribution and response in SOD and RHD2 loss‐of‐function mutants. Treatment of Arabidopsis seedlings with additional sources of ROS (hydrogen peroxide) or an ROS production inhibitor (diphenylene iodonium) induced phenocopies of the mutants studied. Simultaneous application of auxin and ROS increased LR primordia induction, and PIN‐FORMED protein immunolocalization further demonstrated the existing link between auxin and ROS in orchestrating cell division and auxin flux during root development. In Arabidopsis roots, genetic alterations in ROS balance led to defective auxin distribution and growth‐related responses in roots. Exogenous hydrogen peroxide alters the establishment of the endogenous auxin gradient in the root meristem through regulation of PIN‐FORMED polarity, while the simultaneous application of hydrogen peroxide and auxin enhanced LR induction in a dose‐ and position‐dependent manner through activation of cell division. Significance Statement: In Arabidopsis roots, genetic alterations in reactive oxygen species balance led to defective auxin distribution and growth‐related responses in roots. Exogenous hydrogen peroxide altered the establishment of the endogenous auxin gradient in the root meristem through regulation of PIN‐FORMED polarity, while the simultaneous application of hydrogen peroxide and auxin enhanced lateral root induction in a dose‐ and position‐dependent manner through activation of cell division. [ABSTRACT FROM AUTHOR]