Hormonal and epigenetic regulation of root responses to salinity stress

Salinity stress is a major environmental stress affecting crop productivity,and its negative impact on global food security is only going to increase,due to current climate trends.Salinity tolerance was present in wild crop relatives but significantly weakened during domestication.Regaining it back requires a good understanding of molecular mechanisms and traits involved in control of plant ionic and ROS homeostasis.This review summarizes our current knowledge on the role of major plant hormones(auxin,cytokinins,abscisic acid,salicylic acid,and jasmonate)in plants adaptation to soil salinity.We firstly discuss the role of hormones in controlling root tropisms,root growth and architecture(primary root elongation,meristematic activity,lateral root development,and root hairs formation).Hormone-mediated control of uptake and sequestration of key inorganic ions(sodium,potassium,and calcium)is then discussed followed by regulation of cell redox balance and ROS signaling in salt-stressed roots.Finally,the role of epigenetic alterations such as DNA methylation and histone modifications in control of plant ion and ROS homeostasis and signaling is discussed.This data may help develop novel strategies for breeding and cultivating salt-tolerant crops and improving agricultural productivity in saline regions.

Overexpression of the peroxidase gene ZmPRX1 increases maize seedling drought tolerance by promoting root development and lignification

Drought is a main abiotic stress factor hindering plant growth,development,and crop productivity.Therefore,it is crucial to understand the mechanisms by which plants cope with drought stress.Here,the function of the maize peroxidase gene ZmPRX1 in drought stress tolerance was investigated by measurement of its expression in response to drought treatment both in a ZmPRX1 overexpression line and a mutant line.The higher root lignin accumulation and seedling survival rate of the overexpression line than that of the wild type or mutant support a role for ZmPRX1 in maize drought tolerance by regulating root development and lignification.Additionally,yeast one-hybrid,Dule luciferase and ChIP-qPCR assays showed that ZmPRX1 is negatively regulated by a nuclear-localized ZmWRKY86 transcription factor.The gene could potentially be used for breeding of drought-tolerant cultivars.

The glutamate receptor gene GLR3.3:A bridge of calciummediated root development in poplar

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×P.glandulosa)in response to different calcium concentrations and found that low Ca^(2+)(1 mmol·L^(-1))promoted lateral root development,while deficiency(0.1 mmol·L^(-1)Ca^(2+))inhibited lateral root development.Co-expression analysis showed that Ca^(2+)channel glutamate receptors(GLRs)were present in various modules with significance for root development.Two GLR paralogous genes,PagGLR3.3a and Pag GLR3.3b,were mainly expressed in roots and up-regulated under Ca^(2+)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 Ca^(2+)channels mediated root growth and development in trees,but also provides a directive breeding of new poplar species for biofuel and bioenergy production.

Increase in root density induced by coronatine improves maize drought resistance in North China

Drought stress caused by insufficient irrigation or precipitation impairs agricultural production worldwide.In this study,a two-year field experiment was conducted to investigate the effect of coronatine(COR),a functional analog of jasmonic acid(JA),on maize drought resistance.The experiment included two water treatments(rainfed and irrigation),four COR concentrations(mock,0μmol L^(-1);A1,0.1μmol L^(-1);A2,1μmol L^(-1);A3,10μmol L^(-1))and two maize genotypes(Fumin 985(FM985),a drought-resistant cultivar and Xianyu 335(XY335),a drought-sensitive cultivar).Spraying 1μmol L^(-1)COR at seedling stage increased surface root density and size,including root dry matter by 12.6%,projected root area by 19.0%,average root density by 51.9%,and thus root bleeding sap by 28.2%under drought conditions.COR application also increased leaf area and SPAD values,a result attributed to improvement of the root system and increases in abscisic acid(ABA),JA,and salicylic acid(SA)contents.The improvement of leaves and roots laid the foundation for increasing plant height and dry matter accumulation.COR application reduced anthesis and silking interval,increasing kernel number per ear.COR treatment at 1μmol L^(-1)increased the yield of XY335 and FM985 by 7.9%and 11.0%,respectively.Correlation and path analysis showed that grain yields were correlated with root dry weight and projected root area,increasing maize drought resistance mainly via leaf area index and dry matter accumulation.Overall,COR increased maize drought resistance mainly by increasing root dry weight and root area,with 1μmol L-^(-1)COR as an optimal concentration.

Overexpression of the LcPIN2 and LtPIN2 Gene in Arabidopsis thaliana Promotes Root Elongation

The auxin polar transporter,PIN-FORMED 2(PIN2)plays an important role in root development.However,it remains unclear whether PIN2 genes form two Liriodendron species,L.chinense(LcPIN2)and L.tulipifera(LtPIN2),are both involved in root development and whether and to what extent these two genes diverge in function.Here,we cloned and overexpressed LcPIN2 and LtPIN2 in Arabidopsis thaliana wild-type(WT)and Atpin2 mutant.Phylogenetic and sequence analysis showed a small degree of differentiation between these two Liriodendron PIN2 genes.Tissue-specific gene expression analysis indicated that both Liriodendron PIN2 genes were highly expressed in roots,implying a potential role in root development.Finally,heterologous overexpression of LcPIN2 and LtPIN2 in Arabidopsis both significantly increased the root length compared to wild-type and empty vector.Furthermore,the root length defect in Atpin2 was complemented both by LcPIN2 and LtPIN2.However,heterologous overexpression of LcPIN2 and LtPIN2 cannot rescue the defect in root gravitropism of Atpin2 mutants.Taken together,ourfindings unravel PIN2 genes from the magnoliids plant Liriodendron were functionally conserved with AtPIN2 in the dicotyledonous plant Arabidopsis in regard to the regulation of root length,but not root gravitropism.This study also provides a potential target for genetic improvement of the root system in these valuable forest trees Liriodendron.

The ABA synthesis enzyme allele OsNCED2^(T)promotes dryland adaptation in upland rice

Upland rice shows dryland adaptation in the form of a deeper and denser root system and greater drought resistance than its counterpart,irrigated rice.Our previous study revealed a difference in the frequency of the OsNCED2 gene between upland and irrigated populations.A nonsynonymous mutation(C to T,from irrigated to upland rice)may have led to functional variation fixed by artificial selection,but the exact biological function in dryland adaptation is unclear.In this study,transgenic and association analysis indicated that the domesticated fixed mutation caused functional variation in OsNCED2,increasing ABA levels,root development,and drought tolerance in upland rice under dryland conditions.OsNCED2-overexpressing rice showed increased reactive oxygen species-scavenging abilities and transcription levels of many genes functioning in stress response and development that may regulate root development and drought tolerance.OsNCED2^(T)-NILs showed a denser root system and drought resistance,promoting the yield of rice under dryland conditions.OsNCED2^(T)may confer dryland adaptation in upland rice and may find use in breeding dryland-adapted,water-saving rice.

The IDD Transcription Factors:Their Functions in Plant Development and Environmental Response

INDETERMINATE-DOMAIN proteins(IDDs)are a plant-specific transcription factor family characterized by a conserved ID domain with four zinc finger motifs.Previous studies have demonstrated that IDDs coordinate a diversity of physiological processes and functions in plant growth and development,including floral transition,plant architecture,seed and root development,and hormone signaling.In this review,we especially summarized the latest knowledge on the functions and working models of IDD members in Arabidopsis,rice,and maize,particularly focusing on their role in the regulatory network of biotic and abiotic environmental responses,such as gravity,temperature,water,and pathogens.Understanding these mechanisms underlying the function of IDD proteins in these processes is important for improving crop yields by manipulating their activity.Overall,the review offers valuable insights into the functions and mechanisms of IDD proteins in plants,providing a foundation for further research and potential applications in agriculture.

Effect of Polar Auxin Transport on Rice Root Development

Polar auxin transport (PAT) is critical in plant growth and development, especially polar differentiation and pattern formation. Lots of studies have been performed in dicots while relative less in monocots. Using two kinds of PAT inhibitors, 2, 3, 5-triiodobenzoic acid (TIBA) and 9-hydroxyfluorene-9-carboxylic acid (HFCA), it was shown that PAT is important for rice (Oryza sativa L. cv. Zhonghua 11) root development, including elongation of the primary roots, initiation and elongation of lateral roots, and formation of adventitious roots. Inhibition of PAT resulted in the shortened primary roots, less and shortened lateral and adventitious roots. Exogenously supplemented NAA can partially rescue the formation of adventitious roots but not lateral roots, while low concentration of NAA (0.1 mumol/L) could not rescue either of them, suggesting the possible different mechanisms of lateral and adventitious root initiations. Treatment of 30 mumol/L TIBA did not completely inhibit the initiation of lateral roots, and survival capacities of which were demonstrated through cross section experiments revealing the presence of primordial of lateral roots at different stages. Further studies through localized application of PAT inhibitors indicated that auxin flow, transported from coleoptiles to the base, is not only responsible for the auxin contents in stem nodes but also critical for initiation and elongation of adventitious roots.

Differential response of root morphology to potassium deficient stress among rice genotypes varying in potassium efficiency

Disparity in the root morphology of six rice(Oryza sativa L.) genotypes varying in potassium(K) efficiency was studied with three K levels:5 mg/L(low),10 mg/L(moderate) and 40 mg/L(adequate) in hydroponic culture. Morphological parameters included root length,surface area,volume and count of lateral roots,as well as fine(diameter<0.2 mm) and thick(diameter>0.2 mm) roots. The results indicate that the root growth of all genotypes was reduced under low K,but moderate K deficiency increased the root length of the efficient genotypes. At deficient and moderate K levels,all the efficient rice genotypes developed more fine roots(diameter<0.2 mm) than the inefficient ones. Both fine root count and root surface area were found to be the best parameters to portray K stress in rice. In accordance with the root morphology,higher K concentrations were noted in shoots of the efficient genotypes when grown at moderate and deficient K levels,indicating that root morphology parameters are involved in root uptake for K and in the translocation of K up to shoots. K deficiency affected not only the root morphology,but also the root ultra-structure. The roots of high-efficient genotypes had stronger tolerance to K deficient stress for root membrane damage,and could maintain the developed root architecture to adapt to the low K growth medium.

Plant growth and their root development after inoculation of arbuscular mycorrhizal fungi in coal mine subsided areas

Coal mining often cause serious land degradation, soil erosion, and desertification affecting growth of the local vegetation, especially the roots. Arbuscular mycorrhizal fungi （AMF） inoculation is considered a potential biotechnological tool for mined soil remediation because mycorrhizal fungi could improve plant growth environment, especially under adverse conditions due to their good symbiosis. A field experiment was conducted to study the ecological effects of AMF （Funneliformis mosseae, Rhizophagus intraradices） on the growth of Amygdalus pedunculata Pall. and their root development in the regenerated mining subsidence sandy land. The reclamation experiment included four treatments： inoculation of Funneliformis mosseae （F.m）, inoculation of Rhizophagus intraradices （R.i）, combined inoculation of F.m and R.i and non-inoculated treatment. Root mycorrhizal colonization, plant height, crown width, soil moisture, root morphology and certain soil properties were assessed. The results showed that AMF improved the shoot and root growth of Amygdalus pedunculata Pall., and significantly increased root colonization after 1 year of inoculation. Available phosphorus content, activities of phosphatase as well as electrical conductivity in soil rhizosphere of all the three inoculation treatments were higher than that of the non-inoculated treatment. AMF increased the quantity of bacteria and fungi in soil rhizosphere compared with the non-inoculated treatment. Our study indicates that revegetation with AMF inoculum could influence plant growth and root development as well as soil properties, suggesting that AMF inoculation can be effective method for further ecological restoration in coal mine subsided areas.

Effect of CO_2 Elevation on Root Growth and Its Relationship with Indole Acetic Acid and Ethylene in Tomato Seedlings

A hydroponic experiment was carried out to study the effect of elevated carbon dioxide(CO2) on root growth of tomato seedlings.Compared with the control(350 μL L-1),CO2 enrichment(800 μL L-1) significantly increased the dry matter of both shoot and root,the ratio of root to shoot,total root length,root surface area,root diameter,root volume,and root tip numbers,which are important for forming a strong root system.The elevated CO2 treatment also significantly improved root hair development and elongation,thus enhancing nutrient uptake.Increased indole acetic acid concentration in plant tissues and ethylene release in the elevated CO2 treatment might have resulted in root growth enhancement and root hair development and elongation.

Molecular regulatory mechanism of tooth root development

The root is crucial for the physiological function of the tooth, and a healthy root allows an artificial crown to function as required clinically. Tooth crown development has been studied intensively during the last few decades, but root development remains not well understood. Here we review the root development processes, including cell fate determination, induction of odontoblast and cementoblast differentiation, interaction of root epithelium and mesenchyme, and other molecular mechanisms. This review summarizes our current understanding of the signaling cascades and mechanisms involved in root development. It also sets the stage for de novo tooth regeneration.

Pseudomonas sp.TK35-L enhances tobacco root development and growth by inducing HRGPnt3 expression in plant lateral root formation

Rhizosphere colonization is a key requirement for the application of plant growth-promoting rhizobacteria(PGPR)as a bioferilizer.Signaling molecules are often exchanged between PGPR and plants,and genes in plants may respond to the action of PGPR.Here,the luciferase luxAB gene was electrotransformed into Pseudomonas sp.strain TK35,a PGPR with an afinity for tobacco,and the labelled TK35(TK35-L)was used to monitor colonization dynamics in the tobacco rhizosphere and evaluate the effects of colonization on tobacco growth and root development.The transcript levels of the hydroxyproline rich glycoprotein HRGPnt3 gene,a lateral root induction indicator,in tobacco roots were examined by qPCR.The results showed that TK35-L could survive for long periods in the tobacco rhizosphere and colonize new spaces in the tobacco rhizosphere following tobacco root extension,exhibiting significant increases in root development,seedling growth and potassium accumulation in tobacco plants.The upregulation of HRGPnt3 transcription in the inoculated tobacco suggested that TK35-L can promote tobacco root development by upregulating the transcript levels of the HRGPnt3 gene,which promotes tobacco seedling growth.These findings lay a foundation for future studies on the molecular mechanism underlying the plant growth-promoting activities of PGPR.Futhermore,this work provided an ideal potential strain for biofertilizer production.

Root Function in Nutrient Uptake and Soil Water Effect on NO3^- -N and NH4^+ -N Migration

Root function in uptake of nutrients and the effect of soil water on the transfer and distribution of NO3^--N in arable soil were studied using summer maize （Zea mays L. var. Shandan 9） as a testing crop. Results showed that root growth and water supply had a significant effect on NO3^--N transfer and made NO3^--N distributed evenly from bulk soil to rhizosphere soil. Under a natural condition with irrigation, the difference of NO3^--N concentration at different distance points from a maize plant was smaller, while obvious difference of NO3^--N concentration was observed under conditions of limited root growth space without irrigation. Whether root growth space was restricted or not, the content of soil NO3^--N decreased gradually from 10 to 0 cm from the plant, being opposite to the root absorbing area in soils. When root-grown space was limited, changes of NO3^--N concentration at different distances from a plant were similar to that of water content in tendency. Results showed that NO3^--N could be transferred as solute to plant root systems with water uptake by plants. However, the transfer and distribution of NH4^--N were not influenced by root growth and soil water supply, being different to NO3^--N.

Short-term effects of organic amendments on soil fertility and root growth of rubber trees on Hainan Island, China

Rubber[Hevea brasiliensis(Willd.ex A.Juss.)Müll.Arg.]plantations are the largest cultivated forest type in tropical China.Returning organic materials to the soil will help to maintain the quality and growth of rubber trees.Although many studies have demonstrated that organic waste materials can be used to improve soil fertility and structure to promote root growth,few studies have studied the eff ects of organic amendments on soil fertility and root growth in rubber tree plantations.Here,bagasse,coconut husk or biochar were applied with a chemical fertilizer to test their eff ects on soil properties after 6 months and compared with the eff ects of only the chemical fertilizer.Results showed that the soil organic matter content,total nitrogen,available phosphorus and available potassium after the chemical fertilizer(F)treatment were all signifi cantly lower than after the chemical fertilizer+bagasse(Fba),chemical fertilizer+coconut husk(Fco)or chemical fertilizer+biochar(Fbi)(p<0.05).Soil pH in all organic amendments was higher than in the F treatment,but was only signifi cantly higher in the Fbi treatment.In contrast,soil bulk density in the F treatment was signifi cantly higher than in treatments with the organic amendments(p<0.05).When compared with the F treatment,soil root dry mass increased signifi-cantly by 190%,176%and 33%in Fba,Fco and Fbi treatments,respectively(p<0.05).Similar results were found for root activity,number of root tips,root length,root surface area and root volume.Conclusively,the application of bagasse,coconut husk and biochar increased soil fertility and promoted root growth of rubber trees in the short term.However,bagasse and coconut husk were more eff ective than biochar in improving root growth of rubber trees.

Characterization of wavy root 1,an agravitropism allele,reveals the functions of OsPIN2 in fine regulation of auxin transport and distribution and in ABA biosynthesis and response in rice(Oryza sativa L.)

Root system architecture is influenced by gravity.How the root senses gravity and directs its orientation,so-called gravitropism,is not only a fundamental question in plant biology but also theoretically important for genetic improvement of crop root architecture.However,the mechanism has not been elucidated in most crops.We characterized a rice agravitropism allele,wavy root 1(war1),a loss-of-function allele in OsPIN2,which encodes an auxin efflux transporter.With loss of OsPIN2 function,war1 leads to altered root system architecture including wavy root,larger root distribution angle,and shallower root system due to the loss of gravitropic perception in root tips.In the war1 mutant,polar auxin transport was disrupted in the root tip,leading to abnormal auxin levels and disturbed auxin transport and distribution in columella cells.Amyloplast sedimentation,an important process in gravitropic sensing,was also decreased in root tip columella cells.The results indicated that OsPIN2 controls gravitropism by finely regulating auxin transport,distribution and levels,and amyloplast sedimentation in root tips.We identified a novel role of OsPIN2 in regulating ABA biosynthesis and response pathways.Loss of OsPIN2 function in the war1 resulted in increased sensitivity to ABA in seed germination,increased ABA level,changes in ABA-associated genes in roots,and decreased drought tolerance in the seedlings.These results suggest that the auxin transporter OsPIN2 not only modulates auxin transport to control root gravitropism,but also functions in ABA signaling to affect seed germination and root development,probably by mediating crosstalk between auxin and ABA pathways.

Review on Mutation in Lateral Root of Rice

Rice roots include seminal roots, adventitious roots, lateral roots and root hairs, At present, progresses in the research of rice roots have been achieved in many aspects, such as root morphology, root activity, root reaction to various environmental factors as a contribution of root growth and rice yield, the relationship between root growth and stems/leaves/flowers/rice, genetic laws of root characters, etc. However, there are very few researches on lateral root mutant. This paper reviewed progresses of the lateral root mutant of rice from the perspectives of phytomorphology to plant physiology and biochemistry to the gene mapping, consisting of mechanism of developing lateral root of rice, gene cloning and functional analysis of lateral root development, the relationship between auxin and lateral roots, agronomic traits of lateral roots mutant, structure and morphology of root hairs, gravity anomaly of root, redox metabolism and proteomics researches of the mutation in lateral root of rice.

Nanosilver-Promoted Lateral Root Development in Rice is Mediated through Hydrogen Peroxide

Nanosilver(10−9 m)refers to particles comprising 20–15,000 silver atoms,exhibiting high stability and specific surface area.At present,nanosilver has been used in agricultural cultivation and production.This study examined the effects of nanosilver on growth and development of rice root systems.Study results showed that fresh weight of rice belowground organs and root length both increased significantly by 5%and 25%,respectively,after rice radicles were treated with 2 ppm of nanosilver for three days.However,the H_(2)O_(2) level reached its peak at 2 days from treatment,but the activities of the antioxidant enzymes CAT,APX,and GR were inhibited by 2 ppm of nanosilver treatment.The results showed that nanosilver treatment inhibited the antioxidant enzyme activity of rice roots.The treatment of rice radicles with 5μM H_(2)O_(2) promoted root development and the same was observed when nanosilver was used for treatment.Moreover,ascorbic acid(AsA)is a H_(2)O_(2) scavenger and therefore rice root development was inhibited when AsA was added to rice radicles together with either treatment of nanosilver or H_(2)O_(2).In summary,nanosilver treatment of rice radicles promoted root growth and development via the regulation of H_(2)O_(2) and not the O2
−pathway.

The BTB/TAZ domain-containing protein CmBT1-mediated CmANR1 ubiquitination negatively regulates root development in chrysanthemum

Roots are fundamental for plants to adapt to variable environmental conditions.The development of a robust root system is orchestrated by numerous genetic determinants and,among them,the MADS-box gene ANR1 has garnered substantial attention.Prior research has demonstrated that,in chrysanthemum,CmANR1positively regulates root system development.Nevertheless,the upstream regulators involved in the CmANR1-mediated regulation of root development remain unidentified.In this study,we successfully identified bric-a-brac,tramtrack and broad(BTB)and transcription adapter putative zinc finger(TAZ)domain protein CmBT1 as the interacting partner of CmANR1 through a yeasttwo-hybrid(Y2H)screening library.Furthermore,we validated this physical interaction through bimolecular fluorescence complementation and pull-down assays.Functional assays revealed that CmBT1 exerted a negative influence on root development in chrysanthemum.In both in vitro and in vivo assays,it was evident that CmBT1mediated the ubiquitination of CmANR1 through the ubiquitin/26S proteasome pathway.This ubiquitination subsequently led to the degradation of the CmANR1 protein and a reduction in the transcription of CmANR1-targeted gene CmPIN2,which was crucial for root development in chrysanthemum.Genetic analysis suggested that CmBT1 modulated root development,at least in part,by regulating the level of CmANR1 protein.Collectively,these findings shed new light on the regulatory role of CmBT1 in degrading CmANR1 through ubiquitination,thereby repressing the expression of its targeted gene and inhibiting root development in chrysanthemum.

Single-cell network analysis reveals gene expression programs for Arabidopsis root development and metabolism

Single-cell RNA-sequencing datasets of Arabidopsis roots have been generated,but related comprehensive gene co-expression network analyses are lacking.We conducted a single-cell gene co-expression network analysis with publicly available scRNA-seq datasets of Arabidopsis roots using a SingleCellGGM algorithm.The analysis identified 149 gene co-expression modules,which we considered to be gene expression programs(GEPs).By examining their spatiotemporal expression,we identified GEPs specifically expressed in major root cell types along their developmental trajectories.These GEPs define gene programs regulating root cell development at different stages and are enriched with relevant developmental regulators.As examples,a GEP specific for the quiescent center(QC)contains 20 genes regulating QC and stem cell niche homeostasis,and four GEPs are expressed in sieve elements(SEs)from early to late developmental stages,with the early-stage GEP containing 17 known SE developmental regulators.We also identified GEPs for metabolic pathways with cell-type-specific expression,suggesting the existence of cell-type-specific metabolism in roots.Using the GEPs,we discovered and verified a columellaspecific gene,NRL27,as a regulator of the auxin-related root gravitropism response.Our analysis thus systematically reveals GEPs that regulate Arabidopsis root development and metabolism and provides ample resources for root biology studies.