Late Embryogenesis Abundant Gene Family Research Articles

Genome-wide identification of the LEA gene family in Panax ginseng: Evidence for the role of PgLEA2-50 in plant abiotic stress response

Ginseng frequently encounters environmental stress during its growth and development. Late Embryogenesis Abundant (LEA) proteins play a crucial role in combating adversity stress, particularly against abiotic challenges In this study, 107 LEA genes from ginseng, spanning eight subfamilies, were identified, demonstrating significant evolutionary conservation, with the LEA2 subfamily being most prominent. Gene duplication events, primarily segmental duplications, have played a major role in the expansion of the LEA gene family, which has undergone strong purifying selection. PgLEAs were unevenly distributed across 22 chromosomes, with each subfamily featuring unique structural domains and conserved motifs. PgLEAs were expressed in various tissues, exhibiting distinct variations in abundance and tissue specificity. Numerous regulatory cis-elements, related to abiotic stress and hormones, were identified in the promoter region. Additionally, PgLEAs were regulated by a diverse array of abiotic stress-related transcription factors. A total of 35 PgLEAs were differentially expressed following treatments with ABA, GA, and IAA. Twenty-three PgLEAs showed significant but varied responses to drought, extreme temperatures, and salinity stress. The transformation of tobacco with the key gene PgLEA2-50 enhanced osmoregulation and antioxidant levels in transgenic lines, improving their resistance to abiotic stress. This study offers insights into functional gene analysis, focusing on LEA proteins, and establishes a foundational framework for research on ginseng's resilience to abiotic stress.

Overexpression of the late embryonic genesis abundant protein MGL3 improves the drought tolerance of maize (Zea mays L.)

AbstractWith the increasing severity of climate change, drought has become a global issue that seriously restricts the development and production of crops. Maize (Zea mays L.) is one of the major food crops in the world. Therefore, the demand for drought‐tolerant maize varieties has been rapidly increasing in the market. Recent studies have found that late embryogenesis abundant (LEA) proteins are involved in plant responses to various osmotic stresses such as drought and salt stress. Thus, we hypothesized that LEA genes may provide similar stress tolerance abilities in maize. We isolated ZmMGL3 of the LEA gene family and developed transgenic maize plants overexpressing ZmMGL3 using Agrobacterium‐mediated transformation. Then, we conducted physiological and biochemical evaluations of the transgenic maize plants exposed to drought stress. Under drought stress (10% polyethylene glycol 6000), the transgenic maize plants showed improved germination rate, seed vigor, radicle length, root length at the seedling stage, and wilting degree after drought and rewatering compared to the wild‐type plants. The transgenic plants also accumulated more catalase, superoxide dismutase, peroxidase, hydrogen peroxide, and superoxide radicals compared to the wild‐type plants. These results indicate that ZmMGL3 enhances drought resistance in maize plants by reducing the content of reactive oxygen species in the leaves and can be used as a candidate gene for the development of drought‐tolerant maize varieties.

Identification and expression analysis of LEA gene family members in pepper (Capsicum annuum L.).

Pepper (Capsicum annuum L.) is an economically important crop containing capsaicinoids in the seed and placenta, which has various culinary, medical, and industrial applications. Late embryogenesis abundant (LEA) proteins are a large group of hydrophilic proteins participating in the plant stress response and seed development. However, to date there have been no genome-wide analyses of the LEA gene family in pepper. In the present study, 82 LEA genes were identified in the C. annuum genome and classified into nine subfamilies. Most CaLEA genes contain few introns (≤ 2) and are unevenly distributed across 10 chromosomes. Eight pairs of tandem duplication genes and two pairs of segmental duplication genes were identified in the LEA gene family; these duplicated genes were highly conserved and may have performed similar functions during evolution. Expression profile analysis indicated that CaLEA genes exhibited different tissue expression patterns, especially during embryonic development and stress response, particularly in cold stress. Three out of five CaLEA genes showed induced expression upon cold treatment. In summary, we have comprehensively reviewed the LEA gene family in pepper, offering a new perspective on the evolution of this family.

Identification of the Maize LEA Gene Family and Its Relationship with Kernel Dehydration

Maize, the most widely planted and highest yielding of the three major crops in the world, requires the development and breeding of new varieties to accommodate the shift towards mechanized harvesting. However, the moisture content of kernels during harvest poses a significant challenge to mechanized harvesting, leading to seed breakage and increased storage costs. Previous studies highlighted the importance of LEA (Late Embryogenesis Abundant) members in regulating kernel dehydration. In this study, we aimed to gain a better understanding of the relationship between the LEA family and grain dehydration in maize. Through expression pattern analysis of maize, we identified 52 LEA genes (ZmLEAs) distributed across 10 chromosomes, organized into seven subgroups based on phylogenetic analysis, gene structure, and conserved motifs. Evolutionary and selective pressure analysis revealed that the amplification of ZmLEA genes primarily resulted from whole-genome or fragment replication events, with strong purifying selection effects during evolution. Furthermore, the transcriptome data of kernels of two maize inbred lines with varying dehydration rates at different developmental stages showed that 14 ZmLEA genes were expressed differentially in the two inbreds. This suggested that the ZmLEA genes might participate in regulating the kernel dehydration rate (KDR) in maize. Overall, this study enhances our understanding of the ZmLEA family and provides a foundation for further research into its role in regulating genes associated with grain dehydration in maize.

Genome-wide identification, characterization, and expression pattern of the late embryogenesis abundant (LEA) gene family in Juglans regia and its wild relatives J. mandshurica

BackgroundLate Embryogenesis Abundant (LEA) proteins are a class of proteins associated with plant stress resistance. Two Juglans species, Juglans regia and J. mandshurica, are both diploid (2n = 32), monoecious perennial economic tree species with high edible, pharmaceutical, and timber value. The identification, characterization, and expression patterns of LEA proteins in J. regia and its wild relative, J. mandshurica, would not only provide the genetic basis of this gene family, but it would also supply clues for further studies of the evolution and regulating mechanisms of LEA proteins in other tree species.ResultsIn this study, we identified 25 and 20 members of the LEA gene family in Juglans regia and its wild relative, Juglans mandshurica, respectively. The results of phylogenetic analysis showed that the LEA members were divided into eight main subgroups. Predictions of their physicochemical properties showed the variable characteristics of LEA proteins, and the subcellular localization analysis indicated that most LEA proteins are localized in the nucleus. Chromosomal localization analysis and gene replication pattern prediction indicated that WGD is the predominant duplication mode of LEA genes. The results of the comparative analysis indicated a high level of collinearity between the two Juglans species. Analysis of cis-acting elements indicated that LEA genes had a relatively wide range of responses to abiotic stresses and phytohormonal processes, particularly in two phytohormones, methyl jasmonate and abscisic acid. Transcriptome profiling and qRT-PCR experiments showed that JrLEAs are commonly expressed in leaves, green husks, and male and female flowers, and most JmLEAs are more highly expressed in male flowers. We also hypothesized that JrLEAs are involved in the process of anthracnose resistance. Anthracnose-resistant varieties of JrLEAs presented relatively high expression levels at later stages.ConclusionIn this study, we provide a theoretical basis for the functional study of LEA genes in J. regia and J. mandshurica. Analysis of cis-acting elements and gene expression indicated that JrLEAs and JmLEAs play important roles in resistance to biotic stresses in these species.

Genome-wide study and functional characterization elucidates the potential association of late embryogenesis abundant (LEA) genes with lotus seed development

Late embryogenesis abundant (LEA) proteins are extremely hydrophilic proteins imperatively associated with plant growth and development, as well as cell protection from abiotic stress. However, the genome-wide characterization of LEA gene family remains limited, especially in aquatic species such as lotus (Nelumbo spp.). Here, 57 putative LEA genes, including 28 NnLEAs and 29 NlLEAs were identified in the N.nucifera and N.lutea genomes, respectively. A total of 27 homologous LEA gene pairs were identified, indicating high degree of sequence homologies between the two Nelumbo species. Secondary structure prediction indicated high prevalence of alpha (α) helix structure among LEA proteins in the LEA_1, LEA_4, and SMP groups. Screening of putative promoter cis-elements revealed that NnLEA genes were involved in diverse biological processes. Most NnLEA genes were predominantly expressed in the late cotyledons and plumules development stages, suggesting their potential vital roles in lotus seed maturation. In addition, genes co-expressed with NnLEAs were involved in ABA signaling, seed maturation, and development processes. Overall, this study provides new insights for the in-depth understanding of the functions of NnLEA proteins in lotus seed development, and could act as a useful reference for the molecular breeding of seeds with prolonged lifespan.

Genome-Wide Analysis of Late Embryogenesis Abundant Protein Gene Family in Vigna Species and Expression of VrLEA Encoding Genes in Vigna glabrescens Reveal Its Role in Heat Tolerance.

Late embryogenesis abundant (LEA) proteins are identified in many crops for their response and role in adaptation to various abiotic stresses, such as drought, salinity, and temperature. The LEA genes have been studied systematically in several crops but not in Vigna crops. In this study, we reported the first comprehensive analysis of the LEA gene family in three legume species, namely, mung bean (Vigna radiata), adzuki bean (Vigna angularis), and cowpea (Vigna unguiculata), and the cross-species expression of VrLEA genes in a wild tetraploid species, Vigna glabrescens. A total of 201 LEA genes from three Vigna crops were identified harboring the LEA conserved motif. Among these 55, 64, and 82 LEA genes were identified in mung bean, adzuki bean, and cowpea genomes, respectively. These LEA genes were grouped into eight different classes. Our analysis revealed that the cowpea genome comprised all eight classes of LEA genes, whereas the LEA-6 class was absent in the mung bean genome. Similarly, LEA-5 and LEA-6 were absent in the adzuki bean genome. The analysis of LEA genes provides an insight into their structural and functional diversity in the Vigna genome. The genes, such as VrLEA-2, VrLEA-40, VrLEA-47, and VrLEA-55, were significantly upregulated in the heat-tolerant genotype under stress conditions indicating the basis of heat tolerance. The successful amplification and expression of VrLEA genes in V. glabrescens indicated the utility of the developed markers in mung bean improvement. The results of this study increase our understanding of LEA genes and provide robust candidate genes for future functional investigations and a basis for improving heat stress tolerance in Vigna crops.

Genome-wide identification and expression analysis of late embryogenesis abundant protein-encoding genes in rye (Secale cereale L.).

Late embryogenesis abundant (LEA) proteins are members of a large and highly diverse family that play critical roles in protecting cells from abiotic stresses and maintaining plant growth and development. However, the identification and biological function of genes of Secale cereale LEA (ScLEA) have been rarely reported. In this study, we identified 112 ScLEA genes, which can be divided into eight groups and are evenly distributed on all rye chromosomes. Structure analysis revealed that members of the same group tend to be highly conserved. We identified 12 pairs of tandem duplication genes and 19 pairs of segmental duplication genes, which may be an expansion way of LEA gene family. Expression profiling analysis revealed obvious temporal and spatial specificity of ScLEA gene expression, with the highest expression levels observed in grains. According to the qRT-PCR analysis, selected ScLEA genes were regulated by various abiotic stresses, especially PEG treatment, decreased temperature, and blue light. Taken together, our results provide a reference for further functional analysis and potential utilization of the ScLEA genes in improving stress tolerance of crops.

Late embryogenesis abundant (LEA) gene family in Salvia miltiorrhiza: identification, expression analysis, and response to drought stress

ABSTRACT Late embryogenesis abundant (LEA) proteins play important roles in plant defense response to drought stress. However, genome-wide identification of the LEA gene family was not revealed in Salvia miltiorrhiza. In this study, 61 SmLEA genes were identified from S. miltiorrhiza and divided into 7 subfamilies according to their conserved domains and phylogenetic relationships. SmLEA genes contained the LEA conserved motifs and few introns. SmLEA genes of the same subfamilies had similar gene structures and predicted subcellular locations. Our results indicated that the promoters of SmLEA genes contained various cis-acting elements associated with abiotic stress response. In addition, RNA-seq and real-time PCR results suggested that SmLEA genes are specifically expressed in different tissue, and most SmLEA genes can be induced by drought stress. These results provide a valuable foundation for future functional investigations of SmLEA genes and drought stress-resistant breeding of S. miltiorrhiza.

Genome-wide identification and characterization of late embryogenesis abundant protein-encoding gene family in wheat: Evolution and expression profiles during development and stress

Late embryogenesis abundant (LEA) proteins are involved in plant stress responses and osmotic regulation, and they are accumulated in the late embryonic stage. There have been no previous genome-wide analyses of the LEA gene family members in wheat and its close relatives. In this study, 281, 53, 151, 89, 99, and 99 LEA genes were identified in wheat (Triticum aestivum), Triticum urartu, Triticum dicoccoides, Aegilops tauschii, barley, and Brachypodium distachyon, respectively. The wheat LEA gene family (TaLEA genes) was divided into eight subfamilies according to the conserved domains. All TaLEA genes contain very few introns (<3) and they are unevenly distributed on the 21 chromosomes. We identified 39 pairs of tandem duplication genes and 9 pairs of segmental duplication genes in the wheat LEA gene family. This proved that the tandem duplication and segmental duplication played an important role in the expansion of the TaLEA gene family. According to published transcriptome data and qRT-PCR analysis, the TaLEA genes exhibit different tissue expression patterns and they are regulated by various abiotic stresses, especially salt and cold stress. This study provides a comprehensive understanding of the wheat LEA gene family.

Genome-wide identification of and functional insights into the late embryogenesis abundant (LEA) gene family in bread wheat (Triticum aestivum)

Late embryogenesis abundant (LEA) proteins are involved in the responses and adaptation of plants to various abiotic stresses, including dehydration, salinity, high temperature, and cold. Here, we report the first comprehensive survey of the LEA gene family in “Chinese Spring” wheat (Triticum aestivum). A total of 179 TaLEA genes were identified in T. aestivum and classified into eight groups. All TaLEA genes harbored the LEA conserved motif and had few introns. TaLEA genes belonging to the same group exhibited similar gene structures and chromosomal locations. Our results revealed that most TaLEA genes contained abscisic acid (ABA)-responsive elements (ABREs) and various cis-acting elements associated with the stress response in the promoter region and were induced under ABA and abiotic stress treatments. In addition, 8 genes representing each group were introduced into E. coli and yeast to investigate the protective function of TaLEAs under heat and salt stress. TaLEAs enhanced the tolerance of E. coli and yeast to salt and heat, indicating that these proteins have protective functions in host cells under stress conditions. These results increase our understanding of LEA genes and provide robust candidate genes for future functional investigations aimed at improving the stress tolerance of wheat.

Genome-wide identification of the LEA protein gene family in grapevine (Vitis vinifera L.)

The Late Embryogenesis Abundant (LEA) protein gene family is associated with salt and drought stress tolerance in many plant species. However, the number of LEA-encoding genes in the grape (Vitis vinifera) genome has not been previously determined. Here, we conducted a phylogenetic analysis to identify individual members of the LEA gene family in V. vinifera. This analysis identified 60 members of this gene family classified into nine subfamilies (DHN, LEA1, LEA2, LEA3, LEA4, LEA5, LEA6, WHY and SMP) in the grape genome. With 35 members, the LEA2 subfamily constituted the largest subgroup. LEA-encoding genes are non-randomly distributed across most grape chromosomes, suggesting that, similarly to other species, a significant portion of VvLEAs was originated by segmental or tandem duplications. Bioinformatic analyses suggested that the majority of the grape LEA2 proteins is associated with plasma membrane whilst many members of other families are likely to be found in different subcellular locations such as cytoplasm, nucleus and mitochondria. The promoters of these genes contain cis-acting sequence elements associated with stress responses. Expression profiles of a subset of LEA-encoding genes from publically available Affymetrix Vitis vinifera (grape) Genome Array expression data showed that these genes are responsive to salt and dehydration. We also experimentally studied stress-responsive expression patterns of VvDHN1 encoding a dehydrin (DHN) and VvLEA D29L (VvLEA D29Like) encoding a LEA4 protein in the grape cultivar V. vinifera L. cvs. Cabernet Sauvignon (CS) and the rootstock Solonis X riparia 1616 C (1616 C) under salt- and PEG-induced drought stress by qRT-PCR. Both VvDHN1 and VvLEA D29L responded very strongly to these stresses in 1616 C whilst VvLEA D29L was induced only in CS. In response to PEG, VvLEA D29L was induced in both genotypes whilst VvDHN1 showed induction only in CS. Overall, our results suggest the LEA gene family is associated with stress tolerance and manipulation of these genes might provide increased stress tolerance in grapes.

The late embryogenesis abundant gene family in tea plant (Camellia sinensis): Genome-wide characterization and expression analysis in response to cold and dehydration stress

Late embryogenesis abundant (LEA) proteins are a large and highly diverse family of polypeptides that play important roles in plant growth, development and stress responses. At present, LEA gene families have been identified and systematically characterized in many plant species. However, the LEA gene family in tea plant has not been revealed, and the biological functions of the members of this family remain unknown. In this study, 33 CsLEA genes were identified from tea plant via a genome-wide study, and they were clustered into seven groups according to analyses of their phylogenetic relationships, gene structures and protein conserved motifs. In addition, expression analysis revealed that the CsLEA genes were specifically expressed in one or more tissues and significantly induced under cold and dehydration stresses, implying that CsLEA genes play important roles in tea plant growth, development and response to cold and dehydration stresses. Furthermore, a potential transcriptional regulatory network, including DREB/CBF, MYB, bZIP, bHLH, BPC and other transcription factors, is directly associated with the expression of CsLEA genes, which may be ubiquitous and important in the above mentioned processes. This study could help to increase our understanding of CsLEA proteins and their contributions to stress tolerance in tea plant.

Genome-wide identification, structural analysis and new insights into late embryogenesis abundant (LEA) gene family formation pattern in Brassica napus.

Late embryogenesis abundant (LEA) proteins are a diverse and large group of polypeptides that play important roles in desiccation and freezing tolerance in plants. The LEA family has been systematically characterized in some plants but not Brassica napus. In this study, 108 BnLEA genes were identified in the B. napus genome and classified into eight families based on their conserved domains. Protein sequence alignments revealed an abundance of alanine, lysine and glutamic acid residues in BnLEA proteins. The BnLEA gene structure has few introns (<3), and they are distributed unevenly across all 19 chromosomes in B. napus, occurring as gene clusters in chromosomes A9, C2, C4 and C5. More than two-thirds of the BnLEA genes are associated with segmental duplication. Synteny analysis revealed that most LEA genes are conserved, although gene losses or gains were also identified. These results suggest that segmental duplication and whole-genome duplication played a major role in the expansion of the BnLEA gene family. Expression profiles analysis indicated that expression of most BnLEAs was increased in leaves and late stage seeds. This study presents a comprehensive overview of the LEA gene family in B. napus and provides new insights into the formation of this family.

Identification and expression analysis of LEA gene family members in cucumber genome

LEA (late embryogenesis abundant) proteins are firstly discovered in seeds and then identified in vegetative tissues of different plant species. They are mainly regulated under abiotic stress conditions. Although genome wide studies of different gene family members have been performed in cucumber, there is no such a study for LEA genes. We have identified 79 LEA genes in the cucumber genome. Based on phylogenetic analysis, CsLEA genes could be classified into seven groups in which structural motifs are relatively conserved. Tandem duplications play an important role in cucumber genome for LEA gene expansion. Orthologous and chromosomal relationships of CsLEA genes were observed based on comparative mapping analysis with other species. The in silico micro-RNA (miRNA) target analyses indicated that 37 CsLEA genes were targeted by different miRNAs, especially mir854 and mir414 are the most abundant identified ones. Public available RNA-seq data were analyzed for expression analysis of CsLEA genes in different tissues of cucumber. According to genome-wide expression analysis, nine CsLEA genes showed higher expression profiles in all tissues. The expression profiles of ten CsLEA genes in the root and leaf tissues of drought-stressed cucumber were examined using qRT-PCR. Among them, CsLEA-54 induced after stress application in leaf and root tissues and might provide adaptation to drought stress for cucumber. CsLEA-09, CsLEA-32 and CsLEA-57 genes responded to drought after 3 h later and might be considered as early response genes to water limitation. This research could help us to improve understanding of contribution of CsLEAs to drought tolerance in cucumber.

Late Embryogenesis Abundant (LEA) Constitutes a Large and Diverse Family of Proteins Involved in Development and Abiotic Stress Responses in Sweet Orange (Citrus sinensis L. Osb.).

Late Embryogenesis Abundant (LEA) proteins are an ubiquitous group of polypeptides that were first described to accumulate during plant seed dehydration, at the later stages of embryogenesis. Since then they have also been recorded in vegetative plant tissues experiencing water limitation and in anhydrobiotic bacteria and invertebrates and, thereby, correlated with the acquisition of desiccation tolerance. This study provides the first comprehensive study about the LEA gene family in sweet orange (Citrus sinensis L. Osb.), the most important and widely grown fruit crop around the world. A surprisingly high number (72) of genes encoding C. sinensis LEAs (CsLEAs) were identified and classified into seven groups (LEA_1, LEA_2, LEA_3 and LEA_4, LEA_5, DEHYDRIN and SMP) based on their predicted amino acid sequences and also on their phylogenetic relationships with the complete set of Arabidopsis thaliana LEA proteins (AtLEAs). Approximately 60% of the CsLEAs identified in this study belongs to the unusual LEA_2 group of more hydrophobic LEA proteins, while the other LEA groups contained a relatively small number of members typically hydrophilic. A correlation between gene structure and motif composition was observed within each LEA group. Investigation of their chromosomal localizations revealed that the CsLEAs were non-randomly distributed across all nine chromosomes and that 33% of all CsLEAs are segmentally or tandemly duplicated genes. Analysis of the upstream sequences required for transcription revealed the presence of various stress-responsive cis-acting regulatory elements in the promoter regions of CsLEAs, including ABRE, DRE/CRT, MYBS and LTRE. Expression analysis using both RNA-seq data and quantitative real-time RT-PCR (qPCR) revealed that the CsLEA genes are widely expressed in various tissues, and that many genes containing the ABRE promoter sequence are induced by drought, salt and PEG. These results provide a useful reference for further exploration of the CsLEAs functions and applications on crop improvement.

Late Embryogenesis Abundant (LEA) Gene Family in Maize: Identification, Evolution, and Expression Profiles

Late embryogenesis abundant (LEA) proteins are identified as a large and highly diverse group of polypeptides accumulating in response to cellular dehydration in many organisms. However, there are only very limited reports of this protein family in maize until this study. In the present paper, we identified 32 LEA genes in maize. A total of 83 LEA proteins including 51 members in Arabidopsis and 32 putative members in maize were classified into nine groups. Gene organization and motif compositions of the LEA members are highly conserved in each of the groups, indicative of their functional conservation. The predicted ZmLEA genes were non-random distributed across chromosomes, and transposition event and segmental duplication contributed to the expansion of the LEA gene family in maize. Some abiotic stress-responsive cis-elements were also found in the promoters of ZmLEA genes. Microarray expression analyses revealed different accumulation patterns of ZmLEA family members. Moreover, some members of ZmLEAs were regulated under IAA and some abiotic stresses. This study will provide comprehensive information for maize LEA gene family and may pave the way for deciphering their functions in further studies.

Identification and phylogenetic analysis of late embryogenesis abundant proteins family in tomato (Solanum lycopersicum).

This study provided a comparative genomic analysis of the LEA gene family, and these may provide valuable information for their functional investigations in the future. Late embryogenesis abundant (LEA) proteins are a group of proteins that accumulate in response to cellular dehydration in many organisms. Here, we identified 27 LEA genes in tomato. A strong correlation between phylogeny, gene structure, and motif composition was found. The predicted SlLEA genes were non-randomly distributed within their chromosomes, and segmental and tandem duplications were probably important for their expansion. Many cis-elements potentially mediating transcription in response to abiotic stress were also found in the 1,000bp upstream sequence of the promoter region. An additional intragenic recombination played an important role in the evolution of SlLEA genes. Selection analysis also identified some significant site-specific constraints that acted on the evolution of most LEA paralogs. Expression analysis using both microarray data and quantitative real-time PCR indicated that SlLEA genes were widely expressed in various tissues, and that a few members responded to some abiotic stresses. Our study provides useful information on the LEA genes in tomato and will facilitate their further characterization to better understand their functions.

Genome-wide analysis of the LEA (late embryogenesis abundant) protein gene family in Populus trichocarpa

Late embryogenesis abundant (LEA) proteins are a large and highly diverse group of polypeptides that are believed to function in desiccation and freezing tolerance in plants. This report presents a genome-wide analysis of LEA proteins and their encoding genes in Populus trichocarpa. Fifty-three LEA genes were identified from the Populus genome and divided into eight groups. The LEA4 and LEA5 groups were found in green algae and all land plants, whereas the other six groups existed only in land plants, indicating that the LEA family underwent rapid expansion during the early evolution of land plants. A majority of Populus LEA proteins contained repeated motifs that were often specific to a LEA group. Except for PtLEA2-1 and PtLEA2-3, all Populus LEA proteins were highly hydrophilic. Examination of the chromosomal locations of Populus LEA genes revealed that 30 % were arranged in tandem repeats, indicating that tandem duplications significantly contributed to the expansion of this gene family in Populus. Expression patterns of all Populus LEA genes under normal growth conditions and abiotic stress (salinity and drought) were investigated by reverse transcription polymerase chain reaction. Twelve of 53 Populus LEA genes were selectively expressed in a specific tissue and/or in response to a specific treatment. LEA genes also showed extensive divergence in expression patterns, even among those from the same group or gene cluster. The expression profiles revealed that the Populus LEA gene family was not involved systematically in the same regulatory pathway.