Model Legume Plant Research Articles

Genome-wide identification and characterization of filamentation temperature-sensitive H (FtsH) genes and expression analysis in response to multiple stresses in Medicago truncatula.

Filamentation temperature-sensitive H (FtsH) is an AAA+ ATP-dependent protease that plays a vital role in plant environmental adaption and tolerance. However, little is known about the function of the FtsH gene family in the most important legume model plant, Medicago truncatula. To identify and investigate the potential stress adaptation roles of FtsH gene family in M. truncatula, we conducted a series of genome-wide characterization and expression analyses. Totally, twenty MtFtsH genes were identified, which were unevenly distributed across eight chromosomes and classified into six evolution groups based on their phylogenetic relationships, with each group containing similar structures and motifs. Furthermore, MtFtsH genes exhibited a high degree of collinearity and homology with leguminous plants such as alfalfa and soybean. Multiple cis-elements in the upstream region of MtFtsH genes were also identified that responded to light, abiotic stress, and phytohormones. Public RNA-seq data indicated that MtFtsH genes were induced under both salt and drought stresses, and our transcript expression analysis showed that MtFtsH genes of MtFtsH1, MtFtsH2, MtFtsH4, MtFtsH9, and MtFtsH10 were up-regulated after ABA, H2O2, PEG, and NaCl treatments. These results suggest that MtFtsH genes may play a critical role in drought and high salt stress responses and the adaption processes of plants. This study provides a systematic analysis of FtsH gene family in M. truncatula, serving as a valuable molecular theoretical basis for future functional investigations. Our findings also extend the pool of potential candidate genes for the genetic improvement of abiotic stress tolerance in legume crops.

Integrated bioinformatic and physiological analyses reveal the pivotal role of hydrogen sulfide in enhancing low-temperature tolerance in alfalfa.

Hydrogen sulfide (H2 S) is an important gaseous signal molecule that regulates plant growth and stress resistance. However, research on the H2 S synthase (HSase) genes is still limited in the model legume plant Medicago truncatula Gaertn. In the present study, a total of 40 HSase family members were first identified and analyzed in the M. truncatula genome, and these genes distributed across eight chromosomes and were clustered into five groups (I-V) based on their conserved gene structures and protein motifs. Expression analysis revealed that the MtHSase genes were expressed in all the tested abiotic stresses, albeit with expression level differences. This study also showed that H2 S improves low temperature tolerance of alfalfa seedlings by regulating the antioxidant defense system and enhancing photosynthetic capacity. Thus, the study provides new insights into how the H2 S signal regulates tolerance to low-temperature stress and provides the basis for further gene function and detection.

Reorganization of three-dimensional chromatin architecture in Medicago truncatula under phosphorus deficiency.

Emerging evidence reveals that the three-dimensional (3D) chromatin architecture plays a key regulatory role in various biological processes of plants. However, information on the 3D chromatin architecture of the legume model plant Medicago truncatula and its potential roles in the regulation of response to mineral nutrient deficiency are very limited. Using high-resolution chromosome conformation capture sequencing, we identified the 3D genome structure of M. truncatula in terms of A/B compartments, topologically associated domains (TADs) and chromatin loops. The gene density, expressional level, and active histone modification were higher in A compartments than in B compartments. Moreover, we analysed the 3D chromatin architecture reorganization in response to phosphorus (P) deficiency. The intra-chromosomal cis-interaction proportion was increased by P deficiency, and a total of 748 A/B compartment switch regions were detected. In these regions, density changes in H3K4me3 and H3K27ac modifications were associated with expression of P deficiency-responsive genes involved in root system architecture and hormonal responses. Furthermore, these genes enhanced P uptake and mobilization by increasing root surface area and strengthening signal transduction under P deficiency. These findings advance our understanding of the potential roles of 3D chromatin architecture in responses of plants in general, and in particular in M. truncatula, to P deficiency.

Use of halotolerant Bacillus amyloliquefaciens RHF6 as a bio-based strategy for alleviating salinity stress in Lotus japonicus cv Gifu

Halotolerant (HT) bacteria are a group of microorganisms able to thrive in environments with relatively high salt concentrations. HT-microorganisms with plant growth-promoting (PGP) characteristics have been proposed to increase plant tolerance in salty soil. Here, we evaluated the PGP properties at increasing NaCl concentrations of HT-Bacillus strains, previously shown to have beneficial effects under physiological conditions. Most of the isolated showed indole acetic acid and ammonia production and were able to solubilize phosphate and suppress the proliferation of the phytopathogenic fungus Macrophomina phaseolina 2013–1 at high salt concentrations. One of the selected strains, Bacillus amyloliquefaciens RHF6, which retained its beneficial properties up to 400 mM NaCl in vitro, was tested on the legume model plant Lotus japonicus cv Gifu under salt stress. The inoculation with RHF6 significantly improved the survival of plants under high salinity conditions, as reflected in seedling root and shoot growth and total fresh weight (increased by 40%) when compared with non-inoculated plants. The ability of RHF6 to induce a plant antioxidant response, secrete the osmoprotectant proline and reduce ethylene level via the enzymatic ACC deaminase activity indicated this strain as a potentially helpful PGPB for the treatment of degraded soils.

Core clock component MtLUX controls shoot architecture through repression of MtTB1/MtTCP1A in Medicago truncatula

Plants are capable of regulating their shoot architecture in response to diverse internal and external environments. The circadian clock is an adaptive mechanism that integrates information from internal and ambient conditions to help plants cope with recurring environmental fluctuations. Despite the current understanding of plant circadian clock and genetic framework underlying plant shoot architecture, the intricate connection between these two adaptive mechanisms remains largely unclear. In this study, we elucidated how the core clock gene LUX ARRHYTHMO (LUX) regulates shoot architecture in the model legume plant Medicago truncatula. We show that mtlux mutant displays increased main stem height, reduced lateral shoot length, and decreased the number of lateral branches and biomass yield. Gene expression analysis revealed that MtLUX regulated shoot architecture by repressing the expression of strigolactone receptor MtD14 and MtTB1/MtTCP1A, a TCP gene that functions centrally in modulating shoot architecture. In vivo and in vitro experiments showed that MtLUX directly binds to a cis-element in the promoter of MtTB1/MtTCP1A, suggesting that MtLUX regulates branching by rhythmically suppressing MtTB1/MtTCP1A. This work demonstrates the regulatory effect of the circadian clock on shoot architecture, offering a new understanding underlying the genetic basis towards the flexibility of plant shoot architecture.

Identification and characterization of long noncoding RNAs involved in the aluminum stress response in Medicago truncatula via genome-wide analysis.

Numerous studies have shown that plant long noncoding RNAs (lncRNAs) play an important regulatory role in the plant response to environmental stress. However, there are no reports on lncRNAs regulating and enhancing aluminum (Al) stress tolerance in legumes. This study analyzed the role of lncRNAs in response to Al stress in the legume model plant Medicago truncatula. A total of 219.49 Gb clean data were generated: 3,284 lncRNA genes were identified, of which 515 were differentially expressed, and 1,254 new genes were functionally annotated through database alignment. We further predicted and classified putative targets of these lncRNAs and found that they were enriched in biological processes and metabolic pathways such as plant hormone signal transduction, cell wall modification and the tricarboxylic acid (TCA) cycle. Finally, we characterized the functions of 2 Al-activated-malate-transporter-related lncRNAs in yeast. The recombinant plasmids of MSTRG.12506.5 and MSTRG.34338.20 were transformed into yeast, and these yeast exhibited better growth than those carrying empty vectors on medium supplemented with 10 μM AlCl3 and showed that they have biological functions affording Al stress tolerance. These findings suggest that lncRNAs are involved in regulating plant responses to Al stress. Our findings help to understand the role of lncRNAs in the response to Al stress in legumes and provide candidate lncRNAs for further studies.

The mycorrhizal symbiosis alters the plant defence strategy in a model legume plant.

Arbuscular mycorrhizal (AM) symbiosis modulates plant-herbivore interactions. Still, how it shapes the overall plant defence strategy and the mechanisms involved remain unclear. We investigated how AM symbiosis simultaneously modulates plant resistance and tolerance to a shoot herbivore, and explored the underlying mechanisms. Bioassays with Medicago truncatula plants were used to study the effect of the AM fungus Rhizophagus irregularis on plant resistance and tolerance to Spodoptera exigua herbivory. By performing molecular and chemical analyses, we assessed the impact of AM symbiosis on herbivore-triggered phosphate (Pi)- and jasmonate (JA)-related responses. Upon herbivory, AM symbiosis led to an increased leaf Pi content by boosting the mycorrhizal Pi-uptake pathway. This enhanced both plant tolerance and herbivore performance. AM symbiosis counteracted the herbivore-triggered JA burst, reducing plant resistance. To disentangle the role of the mycorrhizal Pi-uptake pathway in the plant's response to herbivory, we used the mutant line ha1-2, impaired in the H+ -ATPase gene HA1, which is essential for Pi-uptake via the mycorrhizal pathway. We found that mycorrhiza-triggered enhancement of herbivore performance was compromised in ha1-2 plants. AM symbiosis thus affects the defence pattern of M. truncatula by altering resistance and tolerance simultaneously. We propose that the mycorrhizal Pi-uptake pathway is involved in the modulation of the plant defence strategy.

Tolerant mechanism of model legume plant Medicago truncatula to drought, salt, and cold stresses.

Legume plants produce one-third of the total yield of primary crops and are important food sources for both humans and animals worldwide. Frequent exposure to abiotic stresses, such as drought, salt, and cold, greatly limits the production of legume crops. Several morphological, physiological, and molecular studies have been conducted to characterize the response and adaptation mechanism to abiotic stresses. The tolerant mechanisms of the model legume plant Medicago truncatula to abiotic stresses have been extensively studied. Although many potential genes and integrated networks underlying the M. truncatula in responding to abiotic stresses have been identified and described, a comprehensive summary of the tolerant mechanism is lacking. In this review, we provide a comprehensive summary of the adaptive mechanism by which M. truncatula responds to drought, salt, and cold stress. We also discuss future research that need to be explored to improve the abiotic tolerance of legume plants.

LjAMT2;2 Promotes Ammonium Nitrogen Transport during Arbuscular Mycorrhizal Fungi Symbiosis in Lotus japonicus.

Arbuscular mycorrhizal fungi (AMF) are important symbiotic microorganisms in soil that engage in symbiotic relationships with legumes, resulting in mycorrhizal symbiosis. Establishment of strong symbiotic relationships between AMF and legumes promotes the absorption of nitrogen by plants. Ammonium nitrogen can be directly utilised by plants following ammonium transport, but there are few reports on ammonium transporters (AMTs) promoting ammonium nitrogen transport during AM symbiosis. Lotus japonicus is a typical legume model plant that hosts AMF. In this study, we analysed the characteristics of the Lotus japonicus ammonium transporter LjAMT2;2, and found that it is a typical ammonium transporter with mycorrhizal-induced and ammonium nitrogen transport-related cis-acting elements in its promoter region. LjAMT2;2 facilitated ammonium transfer in yeast mutant supplement experiments. In the presence of different nitrogen concentrations, the LjAMT2;2 gene was significantly upregulated following inoculation with AMF, and induced by low nitrogen. Overexpression of LjAMT2;2 increased the absorption of ammonium nitrogen, resulting in doubling of nitrogen content in leaves and roots, thus alleviating nitrogen stress and promoting plant growth.

Leaf Development in Medicago truncatula.

Forage yield is largely dependent on leaf development, during which the number of leaves, leaflets, leaf size, and shape are determined. In this mini-review, we briefly summarize recent studies of leaf development in Medicago truncatula, a model plant for legumes, with a focus on factors that could affect biomass of leaves. These include: floral development and related genes, lateral organ boundary genes, auxin biosynthesis, transportation and signaling genes, and WOX related genes.

Integrated Analysis of Coding and Non-coding RNAs Reveals the Molecular Mechanism Underlying Salt Stress Response in Medicago truncatula.

Salt stress is among the most severe abiotic stresses in plants worldwide. Medicago truncatula is a model plant for legumes and analysis of its response to salt stress is helpful for providing valuable insights into breeding. However, few studies have focused on illustrating the whole-transcriptome molecular mechanism underlying salt stress response in Medicago truncatula. Herein, we sampled the leaves of Medicago truncatula treated with water or NaCl and analyzed the characteristics of its coding and non-coding RNAs. We identified a total of 4,693 differentially expressed mRNAs (DEmRNAs), 505 DElncRNAs, 21 DEcircRNAs, and 55 DEmiRNAs. Gene ontology and Kyoto Encyclopedia of Genes and Genomes pathway enrichment analyses revealed that their functions were mostly associated with metabolic processes. We classified the lncRNAs and circRNAs into different types and analyzed their genomic distributions. Furthermore, we predicted the interactions between different RNAs based on the competing endogenous RNA (ceRNA) theory and identified multiple correlation networks, including 27 DEmiRNAs, 43 DEmRNAs, 19 lncRNAs, and 5 DEcircRNAs. In addition, we comprehensively analyzed the candidate DEmRNAs and ceRNAs and found that they were involved in Ca+ signaling, starch and sucrose biosynthesis, phenylpropanoid and lignin metabolism, auxin and jasmonate biosynthesis, and transduction pathways. Our integrated analyses in salt stress response in Medicago truncatula revealed multiple differentially expressed coding and non-coding RNAs, including mRNAs, lncRNAs, circRNAs, and miRNAs, and identified multiple DEmRNA and ceRNA interaction pairs that function in many pathways, providing insights into salt stress response in leguminous plants.

Conservation and Diversity in Gibberellin-Mediated Transcriptional Responses Among Host Plants Forming Distinct Arbuscular Mycorrhizal Morphotypes.

Morphotypes of arbuscular mycorrhizal (AM) symbiosis, Arum, Paris, and Intermediate types, are mainly determined by host plant lineages. It was reported that the phytohormone gibberellin (GA) inhibits the establishment of Arum-type AM symbiosis in legume plants. In contrast, we previously reported that GA promotes the establishment of Paris-type AM symbiosis in Eustoma grandiflorum, while suppressing Arum-type AM symbiosis in a legume model plant, Lotus japonicus. This raises a hitherto unexplored possibility that GA-mediated transcriptional reprogramming during AM symbiosis is different among plant lineages as the AM morphotypes are distinct. Here, our comparative transcriptomics revealed that several symbiosis-related genes were commonly upregulated upon AM fungal colonization in L. japonicus (Arum-type), Daucus carota (Intermediate-type), and E. grandiflorum (Paris-type). Despite of the similarities, the fungal colonization levels and the expression of symbiosis-related genes were suppressed in L. japonicus and D. carota but were promoted in E. grandiflorum in the presence of GA. Moreover, exogenous GA inhibited the expression of genes involved in biosynthetic process of the pre-symbiotic signal component, strigolactone, which resulted in the reduction of its endogenous accumulation in L. japonicus and E. grandiflorum. Additionally, differential regulation of genes involved in sugar metabolism suggested that disaccharides metabolized in AM roots would be different between L. japonicus and D. carota/E. grandiflorum. Therefore, this study uncovered the conserved transcriptional responses during mycorrhization regardless of the distinct AM morphotype. Meanwhile, we also found diverse responses to GA among phylogenetically distant AM host plants.

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MicroRNAs (miRNAs) are a new class of non-protein coding small RNAs that regulate gene expression at the post-transcriptional level in plants and animals. Although thousands of miRNAs were identified in many plant species, only 3 miRNAs have been reported in Lotus Japonicus, a model legume plant. In this study, 80 potential miRNA candidates were identified in 28 ESTs and 52 GSSs of L. japonicus using a homology-based computational analysis. A total of 735 miRNA targets were predicted and some of them encoded transcription factors as well as genes that function in stress response, signal transduction, methylation and others. Quantitative real-time PCR (qRT-PCR) analysis indicated that miR156a, miR160a and miR399a participated in seed germination of L. japonicus. GO and KEGG analysis suggested that the predicted miRNAs might target genes involved in lipid, nitrogen, starch sucrose metabolism and signal transduction.

Medicago ABI3 Splicing Isoforms Regulate the Expression of Different Gene Clusters to Orchestrate Seed Maturation.

Seed maturation comprises important developmental processes, such as seed filling and the acquisition of seed germination capacity, desiccation tolerance, longevity, and dormancy. The molecular regulation of these processes is tightly controlled by the LAFL transcription factors, among which ABSCISIC ACID INSENSITIVE 3 (ABI3) was shown to be involved in most of these seed maturation processes. Here, we studied the ABI3 gene from Medicago truncatula, a model legume plant for seed studies. With the transcriptomes of two loss-of-function Medicago abi3 mutants, we were able to show that many gene classes were impacted by the abi3 mutation at different stages of early, middle, and late seed maturation. We also discovered three MtABI3 expression isoforms, which present contrasting expression patterns during seed development. Moreover, by ectopically expressing these isoforms in Medicago hairy roots generated from the abi3 mutant line background, we showed that each isoform regulated specific gene clusters, suggesting divergent molecular functions. Furthermore, we complemented the Arabidopsis abi3 mutant with each of the three MtABI3 isoforms and concluded that all isoforms were capable of restoring seed viability and desiccation tolerance phenotypes even if not all isoforms complemented the seed color phenotype. Taken together, our results allow a better understanding of the ABI3 network in Medicago during seed development, as well as the discovery of commonly regulated genes from the three MtABI3 isoforms, which can give us new insights into how desiccation tolerance and seed viability are regulated.

Overexpression of MtRAV3 enhances osmotic and salt tolerance and inhibits growth of Medicago truncatula

Related to ABI3/VP1 (RAV) transcription factors play important roles in regulating plant growth and stress tolerance, which have been studied in many plant species, but have remained largely unidentified in legumes. To functionally characterize RAV in legumes, MtRAV3 from legume model plant Medicago truncatula was isolated and its function was investigated by overexpressing MtRAV3 in M. truncatula. Expression analysis demonstrated that MtRAV3 was markedly induced by NaCl and polyethylene glycol (PEG). MtRAV3 overexpression enhanced tolerance of transgenic M. truncatula to mannitol, drought and salt stresses, and induced the expression of adversity-related genes, including MtWRKY76, MtMYB61, cold-acclimation specific protein 31 (MtCAS31), alternative oxidase 1 (MtAOX1) and ethylene response factor 1 (MtERF1). There were lower relative electrolyte leakage and higher chlorophyll content of leaves in the MtRAV3 overexpression plants than in wild type plants under both salt and drought stress. MtRAV3 overexpression M. truncatula were featured by some phenotypes of dwarfing, late flowering, more branches, smaller flower and leaf organs. Further investigations showed that the expression levels of DWARF14 (MtD14), CAROTENOID CLEAVAGE DIOXYGENASES 7 (MtCCD7) and GA3-oxidase1 (MtGA3ox1), which related to dwarf and branch phenotype, were obviously reduced, as well as MtGA3ox1’ (MTR_1g011580), GA20-oxidase1 (MtGA20ox1), MtGA20ox1’ (MTR_1g102070) and GA20-oxidase2 (MtGA20ox2) involved in gibberellins (GAs) pathway. Overall, our results revealed that MtRAV3 exerted an important role in adversity response and plant growth, was a multifunctional gene in M. truncatula, which provided reference for genetic improvement of alfalfa (Medicago sativa).

Genome-wide association study identified candidate genes for seed size and seed composition improvement in M. truncatula

Grain legumes are highly valuable plant species, as they produce seeds with high protein content. Increasing seed protein production and improving seed nutritional quality represent an agronomical challenge in order to promote plant protein consumption of a growing population. In this study, we used the genetic diversity, naturally present in Medicago truncatula, a model plant for legumes, to identify genes/loci regulating seed traits. Indeed, using sequencing data of 162 accessions from the Medicago HAPMAP collection, we performed genome-wide association study for 32 seed traits related to seed size and seed composition such as seed protein content/concentration, sulfur content/concentration. Using different GWAS and postGWAS methods, we identified 79 quantitative trait nucleotides (QTNs) as regulating seed size, 41 QTNs for seed composition related to nitrogen (i.e. storage protein) and sulfur (i.e. sulfur-containing amino acid) concentrations/contents. Furthermore, a strong positive correlation between seed size and protein content was revealed within the selected Medicago HAPMAP collection. In addition, several QTNs showed highly significant associations in different seed phenotypes for further functional validation studies, including one near an RNA-Binding Domain protein, which represents a valuable candidate as central regulator determining both seed size and composition. Finally, our findings in M. truncatula represent valuable resources to be exploitable in many legume crop species such as pea, common bean, and soybean due to its high synteny, which enable rapid transfer of these results into breeding programs and eventually help the improvement of legume grain production.

Sinorhizobium medicae 419 vs S. meliloti 1021: differences in root nodules induced by these two strains on the Medicago truncatula host

Symbiosis between plants of family Fabaceae and bacteria genus Rhizobium is the most widespread interaction, in which atmospheric nitrogen is reduced into ammonia. This highly oxygen-sensitive process is performed by endosymbiotc forms of rhizobia called bacteroids, which occupy newly formed plant organs called root nodules. The goal in this paper was to explore the differences in the (ultra)structure and Reactive Oxygen Species (ROS) localization in two symbiotic interactions between legume model plant Medicago truncatula Gaertn. and bacteria from genus Sinorhizobium. Two bacterial strains—Sinorhizobium meliloti 1021 and Sinorhizobium medicae 419 have the ability of inducing root nodules on the Medicago truncatula, however, such nodules differ in effectiveness of biological nitrogen fixation. We demonstrated that root nodules made of S. medicae 419 [fully effective (F++)] induce a lower number of nodule per plant in comparison to S. meliloti 1021—(partially effective F±). Although the differences in the structure are scant, there are some discrepancies in the ultrastructure. In the meristem of F± nodules, there are Cajal bodies and a lower number of vacuole profiles by larger size in comparison to the F++. Nodules F++ have a thicker interzone and store less starch than F+/, while they have higher esterase activity. Localization of different ROS seems to be comparable in the two type of nodules but interestingly, there is not any Nitric Oxide (NO) in the infection thread of F± root nodules.

Specific tissue proteins 1 and 6 are involved in root biology during normal development and under symbiotic and pathogenic interactions in Medicago truncatula.

ST1 and ST6 are possibly involved in primary and lateral root and symbiotic nodule development, but only ST6 participates in the interaction with hemibiotrophic fungi. Specific tissue (ST) proteins have been shown to be involved in several processes related to plant nutritional status, development, and responses to biotic agents. In particular, ST1 and ST6 are mainly expressed in roots throughout plant development. Here, we analyze where and how the expression of the genes encoding both proteins are modulated in the legume model plant Medicago truncatula in response to the plant developmental program, nodulation induced by a beneficial nitrogen-fixing bacterium (Sinorhizobium meliloti) and the defense response triggered by a pathogenic hemibiotrophic fungus (Fusarium oxysporum). Gene expression results show that ST1 and ST6 participate in the vasculature development of both primary and lateral roots, although only ST6 is related to meristem activity. ST1 and ST6 clearly display different roles in the biotic interactions analyzed, where ST1 is activated in response to a N2-fixing bacterium and ST6 is up-regulated after inoculation with F. oxysporum. The role of ST1 and ST6 in the nodulation process may be related to nodule organogenesis rather than to the establishment of the interaction itself, and an increase in ST6 correlates with the activation of the salicylic acid signaling pathway during the infection and colonization processes. These results further support the role of ST6 in response to hemibiotrophic fungi. This research contributes to the understanding of the complex network that controls root biology and strengthens the idea that ST proteins are involved in several processes such as primary and lateral root development, nodule organogenesis, and the plant-microbe interaction.

Comprehensive identification and characterization of abiotic stress and hormone responsive glycosyl hydrolase family 1 genes in Medicago truncatula

β-glucosidases (BGLUs) hydrolyze the β-D-glycosidic bond with retention of anomeric configuration. BGLUs were associated with many aspects of plant physiological processes, in particular biotic and abiotic stresses through the activation of phytohormones and defense compounds. However, studies on systematic analysis of the stress- or hormone-responsive BGLU genes in plant are still rare. In this study, total 51 BGLU genes of the glycoside hydrolase family 1 were identified in the genome of the model legume plant Medicago truncatula, and they were classified into five distinct clusters. Sequence alignments revealed several conserved and characteristic motifs among these MtBGLU proteins. Analyses of their putative signal peptides and N-glycosylation site suggested that the majority of MtBGLU members have dual targeting to the vacuole and/or chloroplast. Many regulatory elements possibly related with abiotic stresses and phytohormones were identified in MtBGLU genes. Moreover, Microarray and qPCR analyses showed that these MtBGLU genes exhibited distinct expression patterns in various tissues, and in response to different abiotic stress and hormonal treatments. Notably, MtBGLU21, MtBGLU22, MtBGLU28, and MtBGLU30 in cluster I were dramatically activated by NaCl, PEG, IAA, ABA, SA and GA3 treatments. Collectively, our genome-wide characterization, evolutionary analysis, and expression pattern analysis of MtBGLU genes suggested that BGLU genes play crucial roles in response to various abiotic stresses and hormonal cues in M. truncatula.

Calmodulin-Like (CML) Gene Family in Medicago truncatula: Genome-Wide Identification, Characterization and Expression Analysis.

Calcium is an important second messenger in mediating adaptation responses of plants to abiotic and biotic stresses. Calmodulin-like (CML) protein is an important calcium-signaling protein that can sense and decode Ca2+ signal in plants. Medicago truncatula is a model legume plant; however, investigations of MtCML proteins are limited. Using genome analysis and BLAST database searches, fifty MtCML proteins that possess EF-hand motifs were identified. Phylogenetic analysis showed that CML homologs between M. truncatula, Arabidopsis thaliana and Oryza sativa shared close relationships. Gene structure analysis revealed that these MtCML genes contained one to four conserved EF-hand motifs. All MtCMLs are localized to eight chromosomes and underwent gene duplication. In addition, MtCML genes were differentially expressed in different tissues of M. truncatula. Cis-acting elements in promoter region and expression analysis revealed the potential response of MtCML protein to abiotic stress and hormones. The results provide a basis of further functional research on the MtCML gene family and facilitate their potential use for applications in the genetic improvement on M. truncatula in drought, cold and salt stress environments.