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. 2026 Jun 13;26:1116. doi: 10.1186/s12870-026-09230-1

Overexpression of the EfMYB124 gene from Erianthus fulvus enhances tolerance to low temperature and drought in Arabidopsis thaliana

Xibing Rao 1,2, Lilian He 1, Zhenfeng Qian 1, Qingqing Shen 1,3, Zhengying Luo 1, Jun Zhao 2,✉, Fusheng Li 1,✉
PMCID: PMC13326486  PMID: 42288788

Abstract

Background

Environmental factors such as low temperature and water stress often hinder the normal growth of sugarcane (Saccharum spp.), which threatens the supply of raw materials for sugar and biomass provided by sugarcane, resulting in severe economic development losses. Improving the stress resistance of sugarcane has always been one of the important directions in sugarcane breeding. Erianthus fulvus, a wild relative of sugarcane, exhibits remarkable stress tolerance. Therefore, fully exploring the excellent stress-tolerance genes in E. fulvus can provide important candidate genes for improving the stress resistance of sugarcane in future molecular breeding programs.

Results

The evolutionary characteristics of EfMYB124, a gene induced by low temperature and drought, were analysed. Moreover, the gene was successfully transferred into Arabidopsis thaliana, and the degree of damage, physiological responses, and stress-induced gene expression of the transgenic plants under stress were investigated. EfMYB124 is most closely related to At1R-MYB in group A1, and the members of this group have the potential to function in response to adverse stress. Numerous syntenic genes of EfMYB124 were identified in the sugarcane genome. Under drought and low-temperature treatments, the EfMYB124-transgenic plants exhibited relatively mild external damage. Moreover, the activities or contents of three antioxidant enzymes (CAT, POD, and SOD) and Pro in these plants increased rapidly, while the production of MDA decreased. Further analysis revealed that EfMYB124 could specifically regulate the expression of cold stress-related genes (CBF1, CBF3, COR15a, and COR4) and drought stress-related genes (RD29A, P5CS1, SOD1, and CAT2). Additionally, EfMYB124 represses the basal expression levels of key ABA signaling-related genes, including NCED3, ABI5, ABF2, and RD29B.

Conclusions

In summary, this study has initially confirmed that EfMYB124 plays a positive role in regulating plants’ resistance to drought and low temperatures. These key findings lay a solid foundation for us to further analyze the function of EfMYB and study its upstream and downstream interaction relationships. Meanwhile, they provide the necessary scientific reference for the future application of this gene in the genetic improvement of stress tolerance trait breeding in sugarcane.

Supplementary Information

The online version contains supplementary material available at 10.1186/s12870-026-09230-1.

Keywords: Sugarcane, E. fulvus, MYB gene, Drought and cold resistance, Functional verification

Background

Low temperature and drought are important environmental factors that threaten global agricultural production; they not only severely inhibit plant growth but also cause serious damage to plant physiological metabolism and photosynthesis [1–4]. As a result of prolonged evolution, a suite of complex regulatory networks has been established in plants, enabling their adaptation to environmental pressures. When plants respond to adverse stress, they activate the expression of related functional genes so that they can quickly produce adaptive responses in terms of morphology, physiology and metabolism, thus reducing the damage caused by adverse conditions [5–7]. In this process, “star” transcription factor gene families such as NAC, AP2, MYB, and WRKY play a crucial role They can activate downstream genes with specific sequences, thereby regulating plant physiological metabolism and molecular response processes, assisting plants in initiating stress-resistant defence mechanisms, and ultimately enhancing their tolerance to adverse environments [8–12].

The MYB gene family not only has a substantial number of members in most plants but also has an extensive range of biological functions. In most plants, the number of MYB genes exceeds 100 [13–15]. The MYB family of genes consists of four subgroups: R2R3-MYB, the most abundant one; 1R-MYB, the second most common; and R1R2R3-MYB and 4R-MYB, the least common [16]. R1R2R3-MYB is associated mainly with cell cycle regulation. Owing to the very small number of 4R-MYB members, they have received little attention, and their biological functions remain unknown. 1R-MYB and R2R3-MYB are involved in a wide range of gene functions, and there are numerous literature reports on their roles in flowering, cell cycle regulation, metabolite synthesis, response to adverse environments, and more [17, 18]. The homologous expression of the tomato SlMYB102 gene can positively regulate plant cold tolerance through the proline pathway [19]. Following the expression of maize ZmMYB-IF35 in Arabidopsis, transgenic plants were observed to exhibit significantly improved tolerance to oxidative damage under low-temperature stress [20]. Through the regulation of starch breakdown and maltose accumulation, OsMYB30 in rice reinforces cell membrane integrity, ultimately improving cold stress tolerance [21]. After OsMYB55 was expressed in maize, the water deficit damage to the transgenic plants was significantly alleviated, and the plants also presented a relatively high chlorophyll content [22]. Similarly, the TaMYB30-B, TaMYB33, and TaMYB31 genes in wheat have been reported to positively regulate drought tolerance [23–25].

Sugarcane is an important economic crop worldwide; it contributes to the agricultural economic income of more than 100 countries and plays a pivotal role in the supply of sugar raw materials. Concurrently, it supplies approximately 40% of the world’s biofuels, establishing itself as a significant renewable energy crop. Furthermore, the sugar-making process yields numerous valuable byproducts, including cardboard, acetic acid, and rum [26, 27]. However, in the Yunnan and Guangxi regions of China, the growth rate, biological yield, and sugar yield of sugarcane are often affected by low temperature and drought, resulting in significant losses to the economic income of sugarcane farmers. Studies have shown that drought can lead to a 30%-70% reduction in sugarcane yield and a 5% reduction in sucrose formation and sucrose recovery. In addition, low-temperature disasters such as frost in winter and spring can easily and severely affect the quality and buds of sugarcane plants [28]. At present, the widely cultivated sugarcane varieties are almost all hybrid descendants of varieties developed through the “nobilization” approach in the past century. Subsequent generations of backcrossing and hybridisation have resulted in highly polyploid and complex genomes. In addition, owing to the long breeding cycle of sugarcane, coupled with the current scarcity of available excellent innovative parents and low breeding efficiency, it is highly challenging to develop new varieties with high sugar content, high yield and strong stress resistance from scratch. In contrast, improving the genetic traits of sugarcane using modern bioengineering technology has the advantages of short duration, high safety, high efficiency, and strong targeting, making it a feasible approach [29, 30]. Erianthus fulvus is a wild relative of sugarcane that has excellent stress resistance and can grow well even in arid, saline‒alkali, cold and barren areas. It is a very important sugarcane germplasm resource. To date, researchers have assembled the complete genome of diploid sugarcane grass at the chromosome level and constructed a genomic database, which provides favourable support for comprehensively mining the excellent stress resistance genes of sugarcane grass [31–34]. We performed a genome-wide identification and analysis of EfMYB genes in E. fulvus. The results showed that the expression of the 1R-MYB gene EfMYB124 in E. fulvus was significantly and positively induced under low-temperature and drought stresses. In particular, its transcript level was upregulated by 7-fold upon low-temperature treatment, suggesting that this gene possesses crucial potential functions in stress response. Accordingly, EfMYB124 was selected as a key candidate gene for subsequent functional verification [35].

On this basis, we analysed the evolution and characteristics of this gene. Furthermore, we successfully achieved the heterologous overexpression of this gene in Arabidopsis thaliana. Through systematic studies on the performance of transgenic plants exposed to drought and cold stress, we finally revealed the key role of the EfMYB124 gene in this process. Through these analyses, we obtained valuable information about EfMYB124, providing a key basis for future functional analysis, regulatory mechanism research, and molecular breeding applications of the EfMYB124 gene in sugarcane.

Methods

Bioinformatics analysis of EfMYB124

We analysed the phylogenetic relationship between the EfMYB124 protein and At1R-MYB proteins using MEGA 7 (version 7.021) [36, 37]. Additionally, the promoter sequences of the EfMYB124 and At1R-MYB genes were analysed using TBtools (version 2.363) and PlantCARE tools to predict their potential functions [38, 39]. On the other hand, the EfMYB124 protein was submitted to the NCBI Protein Database to search for proteins from other species that are similar to this protein [40]. Subsequent alignment analysis was performed on these homologous proteins.

Synteny analysis

The gene sequence of EfMYB124 was submitted to the Synteny Block of the Saccharum Genome Database to conduct synteny analysis of the EfMYB124 gene in the genomes of Saccharum spontaneum AP85-441 (wild sugarcane species) and sugarcane cultivars (XTT22, R570, and ZZ1) [41–43]. In addition, we constructed a gene sequence information map of these collinear genes to analyze their characteristics.

Plant material

The Columbia ecotype of wild-type A. thaliana was used. First, the seeds were vernalized (4 °C, 3 days), and then all sown in nutrient soil pots with dimensions of 10 × 8 × 10 cm, followed by incubation in a constant temperature incubator at a maintained temperature of 23 °C. The ambient air humidity was set at approximately 60%, the light intensity at 150 µmol/m² ·s, and the daily light duration at 16 h.

Construction of an EfMYB124 overexpression vector

The carrier skeleton used in this study was PU1301-FLAG, preserved by the project team laboratory. First, two single restriction sites, the BamHI and KpnI sites, on the vector backbone were selected for double digestion (overnight at 37 °C). We subsequently further purified the concentration of the vector fragment using a kit. (The kits, gene primers, and PCR systems used in this study are detailed in Table S3; the same applies below). The homologous recombination primers for the EfMYB124 gene were designed by the CE Design V1.04 tool. Using the previously preserved EfMYB124 cloning vector as a template, the homologous arms of EfMYB124 with BamHI and KpnI restriction enzyme sites were amplified by PCR. The purified and linearised PU1301-FLAG vector and the target gene fragment were further connected by homologous recombination using a homologous recombination kit to construct the PU1301-FLAG-EfMYB124 overexpression vector. Finally, the ligation product was identified and subjected to sequencing analysis; after confirming no errors, it was transformed into Agrobacterium GV3101 for subsequent use.

Production of transgenic plants

In this study, EfMYB124 was transferred into A. thaliana by the flower dipping method [44]. First, the positive bacterial solution identified in the previous step was added to a new YEP medium for activation. When the OD600 ranged from 0.8 to 1.0, the bacteria were collected by centrifugation at 4 °C. After 5% sucrose solution was added to resuspend the bacterial solution, the final OD600 was 0.8. Five-week-old A. thaliana plants were used as the transformation recipient material, after which their pods and exposed white flowers were removed. The A. thaliana inflorescence was immersed in the immersion solution for 1–2 min, and then, the A. thaliana was covered with a preservative film for moistening and placed in a dark environment for 24 h. Following infection, the plants were returned to the light incubator for standard growth conditions. One week later, a secondary infection was performed to increase the transformation efficiency.

Following transformation, T0 generation A. thaliana seeds were collected in a sterile 2 mL centrifuge tube, subjected to five washes with sterile water, and then transferred to a new tube of the same type. Subsequently, the seeds were surface-sterilized with 75% ethanol and 5% NaClO solution, rinsed with sterile water, and then air-dried in sterile petri dishes. Seeds were sown evenly on solid MS medium for cultivation, and during this process, positive plants were initially screened using 30 mg/L hygromycin. Genomic DNA was extracted from the leaves of resistant plants. Positive T₁ lines were identified by PCR, and seeds of the T₂ generation were harvested from individual plants. The T₂ seeds were sown on selective medium. Lines exhibiting a 3:1 resistance segregation ratio were individually cultivated and selfed to collect T₃ seeds. Following hygromycin screening, lines with no resistance segregation in their progeny were identified as stably inherited homozygous transgenic T₃ lines.

Assessment of cold and drought tolerance in transgenic A. thaliana

First, the homozygous lines were transplanted into small square pots, with five plants cultivated per pot. After they reached 4 weeks of age, they were used for cold and drought tolerance identification analyses.

In the case of the low-temperature stress treatment, the wild-type (WT) and three overexpression (OE) lines were exposed to -4 °C and 60–70% humidity for 24 h, followed by a recovery phase under standard conditions at 23 °C. Changes in the external phenotypes of A. thaliana were observed on the 3rd and 12th days after recovery. On the other hand, A. thaliana plants were placed in an incubator maintained at a constant temperature of 4 °C. Three stress durations were established: the CK group at 0 h and the treatment groups at 24 h and 72 h. Samples were collected once the respective stress durations were reached. The relevant kits were used to analyse changes in three antioxidant enzymes (CAT, SOD, POD), Pro, and other indicators. In addition, we extracted RNA from these samples and reverse-transcribed it into cDNA. The expression levels of cold-responsive genes (AtCBF3, AtCOR4, AtCOR15a, and AtCBF1) were analysed using a qPCR kit on an ABI 7500 instrument, with the Actin gene serving as the reference gene. We used the 2-ΔΔCt method for the gene expression analysis in all our experiments [45–47].

Drought stress treatment was performed as follows: Robust Arabidopsis plants that were four weeks old were placed in an open-air environment, and then watering was stopped for two weeks to induce drought stress (soil water content: 10 ± 5%). When the specified stress duration is reached, analyze the damage status of the plants. Next, we detected 5 physiological indicators (the types and detection methods of the indicators were the same as above) and analysed their changes under the treatments. To investigate the drought regulatory pathway involving EfMYB124, we analyzed the expression of genes related to antioxidant defense and osmotic regulation, including RD29A, P5CS1, SOD1, and CAT2, as well as representative genes of the ABA signaling pathway, such as RD29B, ABI5, ABF2, and NCED3. In this step, GAPDH was used as a reference gene for normalisation [48–49].

Data analysis

In this study, we designed at least 3 independent replicate experiments for all tests to ensure the reliability of the data. Additionally, all the data in the figures and tables are presented as the average values of three measurements. Data collation was conducted using Excel 2019 software, and one-way analysis of variance was performed with DPS v7.05 software. All figures were generated using Adobe Illustrator 2019 and GraphPad Prism 8.4.2.

Results

Bioinformatics analysis of EfMYB124

When a phylogenetic tree of EfMYB124 and At1R-MYB was constructed, the tree consisted of four groups in total. EfMYB124 clustered in Group A1, along with 11 other At1R-MYB proteins (Fig. 1a). We analyzed the composition of cis-acting elements in the promoters of Group A1 genes. The results revealed that EfMYB124 harbors a specific combination of multiple abiotic stress-responsive elements rarely present in most AtMYB genes. Specifically, its promoter is simultaneously enriched with elements associated with drought induction, low-temperature response, methyl jasmonate responsiveness, abscisic acid response, salicylic acid response, as well as stress defense-related elements. These findings suggest that EfMYB124 possesses the molecular characteristics conferring broad-spectrum stress tolerance. (Fig. 1b). Through sequence alignment in the protein database, 10 proteins with the highest similarity to the EfMYB124 protein were retrieved from other species (Fig. 2). Among them, EfMYB124 shares the highest sequence similarity with MfMYB, followed by ZmMYB. The core MYB domains and motifs of the EfMYB124 protein are highly conserved among these species, whereas its C-terminal regulatory region exhibits species-specific sequence characteristics unique to E. fulvus.

Fig. 1.

Fig. 1

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Phylogenetic tree and cis-acting elements of EfMYB124 and At1R-MYB. a Phylogenetic tree. At1R-MYB is labelled in black font, and EfMYB124 is labelled in red font. b Cis-acting elements in the promoters of genes in Group A1. Rectangular boxes of different colours represent different elements

Fig. 2.

Fig. 2

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Results of multiple sequence alignment. a Phylogenetic relationships among different MYB124 proteins. b The protein sequences of MYB124 from different species. Sequence identity of 100% is marked in black, > 75% in pink, > 50% in cyan, and > 33% in yellow

Synteny analysis of EfMYB124

The synteny analysis results revealed that EfMYB124 had only 3 syntenic genes in the wild sugarcane species S. spontaneum, namely, Sspon.008D0004170, Sspon.008C0005100, and Sspon.008B0004820 (Fig. 3). In contrast, the sugarcane hybrid XTT22 (XTT5 × 69–463) had the most, with 11, followed by the sugarcane hybrids R570 ༈H32-8560 × R445༉ and ZZ1 ༈XTT25 × YZ89-7༉, with 6 and 8, respectively (Fig S1). Among them, the EfMYB124 syntenic genes are predominantly distributed on the homologous chromosomes Chr4 and Chr10 of sugarcane hybrids, whereas those in S. spontaneum are localized on Chr8. These findings indicate that the EfMYB124 gene has undergone different degrees of expansion in sugarcane and its related species and that this expansion is particularly significant in cultivated hybrids. We further analysed the structural characteristics of these syntenic genes and reported that they share high similarity in features such as introns, exons, and coding sequences (Fig. 4). In addition, noticeable variations were observed in intron length and UTR regions. The differences in intron length and the diversity of UTR regions imply that functional divergence may have occurred at the transcriptional regulation level.

Fig. 3.

Fig. 3

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Synteny analysis results. The syntenic regions between the two genomes are represented by grey‒white lines, whereas the yellow lines indicate the syntenic genes of EfMYB124

Fig. 4.

Fig. 4

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Structural characteristics of the EfMYB124 syntenic gene. The black lines represent introns

Analysis of overexpression vector construction

In this study, the plant overexpression vector Pu1301-FLAG-EfMYB124 was successfully constructed using double digestion and homologous recombination techniques. The key vector map is shown in Fig. 5; it contains two promoters, Ubi and 35 S, that regulate the expression of EfMYB124 and Hygr, respectively. The total size of the vector is 15,180 bp, with the EfMYB124 insert being 759 bp. The sequencing results confirmed the accuracy of the gene sequence, and no mutations were detected.

Fig. 5.

Fig. 5

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Structural diagram of the EfMYB124 overexpression vector

Identification of transgenic plants

After subsequent cultivation and screening, 8 hygromycin-resistant plants were ultimately retained. PCR detection of genomic DNA from regenerated plants using Hygr gene-specific primers revealed that 5 out of 8 plants were positive (Fig S2). After verification by qPCR, high expression of the target gene was detected in 3 Arabidopsis plants, successfully obtaining correctly transformed plants (Fig. 6).

Fig. 6.

Fig. 6

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Quantitative analysis of EfMYB124. In the figure, * (p < 0.05), and ** (p < 0.01). All histograms were generated from three biologically repeated datasets. The same applies below

Analysis of the cold tolerance function of EfMYB124

To investigate whether the expression of EfMYB124 can truly increase the resistance of plants to low temperatures, we analysed changes in their growth status following exposure to -4 °C stress. All the plants clearly exhibited vigorous growth before the cold treatment (Fig. 7a). Following exposure to low-temperature stress, the external morphology of individual plants markedly changed, with particularly pronounced differences observed between WT and OE plants. On the 3rd day of recovery, the WT plants showed extensive wilting and yellowing of leaves with severe damage, whereas the three OE lines displayed only minor wilting with noticeably less damage. It is clearly observable that by day 12, all the WT plants were unable to produce new leaves and perished entirely. In contrast, more than 80% of the OE plants resumed healthy growth, indicating robust cold tolerance (Fig. 7b).

Fig. 7.

Fig. 7

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Phenotype of plant cold tolerance. a Phenotypes of OE4, OE8 and OE10 plants under low-temperature stress. b Mortality rates of WT and OE plants following low-temperature stress. The scale bar corresponds to 10 cm

We detected changes in 5 key physiological indicators to determine their physiological adaptation characteristics under cold stress (Fig. 8). When no low-temperature stress was applied, the activities and contents of Pro, CAT, SOD, MDA, and POD in all the plants were similar and did not significantly differ. After the stress was applied (4 °C), the levels of Pro, CAT, POD, MDA and SOD increased in all the A. thaliana lines. Notably, the activity or content of SOD, POD, and Pro in most OE plants was much greater than that in WT plants at the 72 h time point. Compared with the WT plants, the OE8 lines demonstrated a marked increase in CAT activity at both the 24-hour and 72-hour time points. Although not significantly different, compared with the WT plants, the OE4 and OE10 plants also exhibited higher CAT activity. In addition, OE plants exhibited an enhanced capacity for MDA scavenging. Although MDA content increased in all plant lines after stress treatment, OE lines maintained a relatively lower MDA level compared with the WT.

Fig. 8.

Fig. 8

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Physiological response of the plants. a–e represent the activity or content of SOD, POD, CAT, Pro, and MDA in sequence; the same applies below

We performed a quantitative analysis of CBF1, COR15a, COR4, and CBF3 to investigate the regulatory role of EfMYB124 in this process (Fig. 9). The results clearly show that although these genes were upregulated in all the plants after stress treatment, there were significant differences among the plants, with the upregulation range and fold change in the OE being far greater. Notably, at both the 24 h and 72 h treatment time points, the expression levels of CBF1 and CBF3 were significantly greater in all the transgenic plants. On the other hand, when the expression patterns of COR4 and COR15a were compared across different plant lines, the expression levels of these genes in OE plants (excluding OE10) were significantly greater than those in WT plants, which further highlights the unique regulatory effect of EfMYB124 overexpression on these downstream genes.

Fig. 9.

Fig. 9

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qPCR Analysis of CBF1, CBF3, COR4, and COR15a

Analysis of the drought tolerance function of EfMYB124

We analysed the growth status of plants after 14 days of stress treatment to verify whether the expression of EfMYB124 can effectively increase plant drought tolerance. In the absence of stress conditions, all the A. thaliana plants exhibited robust growth with no discernible differences (Fig. 10). After 14 days of drought stress, the WT plants displayed severe wilting, drying, and lodging of leaves, whereas the three OE lines showed only slight wilting. Upon rewatering, the OE lines rapidly resumed growth with favourable recovery states, whereas the WT plants recovered slowly and exhibited poor growth performance.

Fig. 10.

Fig. 10

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Growth alterations in plants under drought stress

To clarify the physiological response characteristics of plants under drought treatment (14 days), we also determined physiological and biochemical indices, including the activity of SOD, CAT, POD, Pro, and MDA (Fig. 11). Under normal conditions, four of these indices (except SOD) remained at low levels across all the plants, with no significant differences observed. Consistent with the findings of previous studies, after stress application, the activities or contents of these indicators increased in all the plants, but the magnitude of the increase was greater in the OE plants. Among these findings, the CAT activity significantly increased in all the transgenic plants. Furthermore, the activity or content of SOD (in OE8), POD (in OE4 and OE10), and Pro (in OE10) in some OE plants was also significantly greater than that in WT plants. Additionally, compared with that of the WT control, the MDA content was consistently lower across all the OE lines.

Fig. 11.

Fig. 11

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Physiological response of the plants

We analysed the gene regulatory mechanism through which EfMYB124 effectively increases plant drought tolerance by examining the expression of four key drought tolerance-related genes (RD29A, P5CS1, SOD1, and CAT2) (Fig. 12). After 14 days of drought stress, the upregulation of these genes was consistently more pronounced in all the OE lines than in the WT plants. Specifically, the expression levels of RD29A and SOD1 were significantly greater in all three OE lines than in the WT plants, whereas P5CS1 and CAT2 expression was significantly greater in OE4 and OE10 than in WT. Another intriguing finding is that under non-stressed basal conditions, EfMYB124 suppresses and downregulates the expression of NCED3, ABI5, ABF2, and RD29B (Fig S3). This suggests that the gene actively represses endogenous ABA biosynthesis and basal activation of the ABA signaling pathway under homeostatic conditions. These findings demonstrate that the pronounced drought resistance of transgenic EfMYB124-overexpressing lines is closely associated with the unique regulatory functions of the EfMYB124 transcription factor. EfMYB124 specifically binds to the promoters of genes involved in antioxidant defense and osmotic adjustment, and modulates their transcription. This regulatory mechanism ultimately constructs a sophisticated molecular network to reinforce plant tolerance to drought stress.

Fig. 12.

Fig. 12

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Changes in RD29A, P5CS1, SOD1 and CAT2 expression

Discussion

In studies on the regulation of stress responses in many plant species, we can observe that MYB genes play a key role. Well-characterised examples include staple crops such as rice and wheat, as well as cash crops such as soybeans and cotton [50–55]. E. fulvus has drawn attention because of its excellent stress tolerance, yet research on MYB genes in this species remains scarce. Based on the research results of the previous stage, EfMYB124 has attracted our attention. Therefore, we conducted a more systematic investigation of this gene in the present study. This gene has potential functions in regulating plant drought and low-temperature stress, and the protein it encodes exhibits high conservation across multiple species, suggesting that these species evolved from a common ancestor. In addition, multiple orthologs of the EfMYB124 gene were identified in the sugarcane genome, with a greater number present in cultivated sugarcane varieties than in their closely related wild species. It is hypothesized that the MYB family is a gene family that has undergone significant expansion and retention in the context of the complex sugarcane genome. This expansion is thought to occur primarily through ancestral polyploidization events or large-segment duplications, rather than through tandem duplications of individual genes. Furthermore, this expansion likely provides additional genetic material and evolutionary potential for the regulation of sugarcane’s adaptability (such as stress resistance) or important agronomic traits. Differences in copy numbers among different varieties may be associated with their respective unique trait performances. To further verify the function of this gene, we overexpressed it in A. thaliana. After low-temperature treatment, the WT plants clearly experienced severe freezing damage followed by complete death, whereas the OE plants experienced only slight injury, rapid regeneration, and strong cold tolerance. The severity of the drought symptoms was markedly more pronounced in the WT plants, which experienced acute dehydration, than in the OE plants, which experienced only slight wilting. Collectively, our findings indicate that EfMYB124 plays a key role in the stress response network of A. thaliana, markedly improving its resilience to both low temperature and water deficit. Previous studies have reported that MYB, Expansins, WRKY and other genes have two or more abiotic stress tolerances at the same time, which was further confirmed in this study [56].

In general, plants’ tolerance to abiotic stress can be reflected in their physiological adaptability. The cell membrane is a critical protective barrier. When plants are exposed to adverse conditions that disrupt their normal physiological metabolism, a series of oxidative stress responses are triggered in plant cells, leading to the excessive accumulation of ROS and various harmful free radicals in vivo. As stress intensifies, membrane lipid peroxidation, membrane substance leakage, and harmful MDA production increase, thereby disrupting cell structure and causing physiological dysfunction [57, 58]. To adapt to these adverse conditions, plants can develop specific physiological responses to mitigate damage. Among these, the adaptive regulation of the antioxidant enzyme system is an important physiological mechanism through which plants resist abiotic stress [59]. For instance, overexpression of MbWRKY40 can increase the activity of antioxidant enzymes under low-temperature stress, thereby effectively preventing further oxidative damage to the cell membrane [60]. To investigate the physiological basis for enhanced stress resistance in transgenic A. thaliana, we analysed the poststress responses of three key antioxidant enzymes and MDA. Under stress, the activity of antioxidant enzymes in OE plants significantly increased, whereas the MDA content relatively decreased, indicating weakened membrane lipid peroxidation. Pro, as an important osmotic regulator and protective agent in plant cells, can modulate cellular osmotic concentration and prevent excessive cellular dehydration [61]. Under both stress treatments, all OE plants presented significantly greater Pro accumulation, and this finding is generally consistent with previous research results [62]. These phenomena imply that compared with WT plants, the expression of the EfMYB124 gene enables transgenic plants to more effectively scavenge ROS, mitigate membrane damage, and balance metabolic processes, thereby allowing them to better withstand harsh environments.

It has been reported that the cold response pathway in plants involves multiple genes and the combined action of several signalling pathways. ICE-CBF/DREB-COR is among the most frequently reported core regulatory networks [63]. CBF genes are rapidly expressed in the cold response pathway, and transgenic A. thaliana plants expressing StCBF1 and StCBF4 exhibit increased tolerance to low temperature [64]. The cold-responsive gene COR, as a downstream gene in the cold response cascade regulatory pathway, can be activated by CBF genes for transcription. This process plays a crucial role in enhancing and regulating the cold tolerance of plants. Specifically, these CBF genes exert their regulatory effects by directly recognising and binding to the conserved CRT/DRE located in the promoter region of target COR genes, thereby initiating the transcription and expression of COR genes to help plants cope with cold stress [65]. Experimental evidence confirms that certain MYB genes act as key transcriptional activators capable of inducing the expression of critical cold-response genes, including CBF and COR. For example, overexpression of MbMYBC1 improved low-temperature tolerance in transgenic A. thaliana, with the upregulation of genes such as COR47, ERD10B, and DREB1A in the former [66]. Similarly, overexpression of the wild strawberry 1R-MYB gene FvMYB114 enhanced cold stress tolerance in transgenic A. thaliana, and FvMYB114 was shown to regulate the expression of CBF1, CBF3, and COR47 [67]. Similarly, Li et al. analysed changes in the expression of CBF1, CBF2, COR15a, and COR78 in FvMYB82-transgenic A. thaliana and reported that their expression significantly increased under low-temperature stress [68]. In summary, the above reports demonstrate that these CBF and COR genes are key genes involved in the response of plants to low temperatures and play crucial roles in resistance to cold damage. To explore this regulatory mechanism, we analysed the changes in the expression of COR15a, COR47, CBF1, and CBF3 and reported that their expression is regulated by EfMYB124. On the basis of these findings, we hypothesise that the EfMYB124 gene may similarly regulate plant cold responses through the canonical CBF signalling pathway. Moreover, in this regulatory process, the interaction of the same type of genes may be involved.

The ABA-dependent signaling pathway is one of the plant drought response pathways, which has been demonstrated in studies on various plants and gene families. Both drought treatment and exogenous ABA treatment can induce the expression of AtMYB44. The overexpression of this gene can effectively alleviate water loss in transgenic plants under drought conditions. Subsequent studies have shown that this function is achieved by AtMYB44, which downregulates the expression of PP2C, a negative regulator of the ABA signalling pathway [69]. Similarly, the expression of the AtMYB15 gene can also be induced by ABA and drought treatments. AtMYB15 effectively promotes the accumulation of ABA in plant cells and increases the efficiency of ABA signal transduction, thereby enhancing the ABA-mediated drought response pathway and helping plants better cope with drought stress [70]. Although drought stress strongly induces the expression of the BcMYB1 gene, it cannot be induced by exogenous ABA treatment, indicating that BcMYB1 may play a role in drought regulation independent of the ABA signalling pathway [71]. Both PEG6000 and MeJA served as strong inducers of the IbMYB116 gene in sweet potato, triggering a substantial increase in its transcript level. Further analysis revealed that in plants overexpressing IbMYB116, genes related to the synthesis of this hormone are regulated by IbMYB116, suggesting that MYB family genes may also regulate drought responses through other signalling pathways [72]. Our analysis results showed that key genes such as P5CS1, SOD1, and CAT2 were all regulated by EfMYB124. This phenomenon was observed in all OE plants, indicating that the enhanced drought tolerance of OE plants is attributed to the regulatory effect of this gene. However, our previous studies have demonstrated that the expression of EfMYB124 is not induced by ABA. In the present study, we further found that the basal expression levels of the ABA synthesis key gene NCED3, the core genes of the ABA signaling pathway (ABI5 and ABF2), and the ABA-dependent drought marker gene RD29B were significantly inhibited under normal growth conditions. We speculate that EfMYB124 modulates drought resistance in A. thaliana mainly via an ABA-independent signaling pathway, rather than by promoting endogenous ABA accumulation or activating the ABA signaling pathway. We plan to further explore its specific mechanism of action in subsequent research. This study preliminarily confirms the role of the EfMYB124 gene in regulating cold and drought tolerance in A. thaliana. These findings pave the way for future research and application of this gene in sugarcane.

Conclusions

In this study, an overexpression vector carrying the low-temperature and drought-induced gene EfMYB124 was constructed and introduced into A. thaliana to investigate its role under low-temperature and drought stress. EfMYB124 promotes physiological adaptive responses and regulates the expression of stress-related genes, playing a positive regulatory role in enhancing low-temperature and drought tolerance in A. thaliana. This is the first report on the functional study of MYB genes in E. fulvus. These key findings lay a solid foundation for us to further analyze the function of EfMYB and study its upstream and downstream interaction relationships. Meanwhile, they provide the necessary scientific reference for the future application of this gene in the genetic improvement of stress tolerance trait breeding in sugarcane.

Supplementary Information

Supplementary Material 1 (12.9KB, xlsx)
Supplementary Material 2 (32.8KB, xlsx)
Supplementary Material 3 (938.8KB, pdf)
Supplementary Material 4 (838.5KB, pdf)
Supplementary Material 5 (148.1KB, pdf)
Supplementary Material 6 (27KB, xlsx)
Supplementary Material 7 (17.1KB, xlsx)

Acknowledgements

We acknowledge American Journal Experts, LLC for their language editing services for the manuscript.

Authors’ contributions

F.S.L., J.Z. and X.B.R. conceived and designed the research; X.B.R. performed the experiments and drafted the manuscript; F.S.L. and J.Z. revised the manuscript; L.L.H., Z.F.Q., Q.Q.S. and Z.Y.L. analysed the data and provided technical assistance. All the authors reviewed and approved the final manuscript.

Funding

This work was supported by the Major Science and Technology Project of Yunnan Province (202202AE090021), the National Natural Science Foundation of China (31960451) and the Guangxi Key Laboratory of Sugarcane Genetic Improvement/Sugarcane Research Institute, Guangxi Academy of Agricultural Sciences (21-238-16-k-02-06).

Data availability

The relevant data can be found in the main text and supplementary materials.

Declarations

Ethics approval and consent to participate

Not applicable.

Consent for publication

Not applicable.

Competing interests

The authors declare no competing interests.

Footnotes

Publisher’s Note

Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Contributor Information

Jun Zhao, Email: junzhao_ky@126.com.

Fusheng Li, Email: 1990017@ynau.edu.cn.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

Supplementary Material 1 (12.9KB, xlsx)
Supplementary Material 2 (32.8KB, xlsx)
Supplementary Material 3 (938.8KB, pdf)
Supplementary Material 4 (838.5KB, pdf)
Supplementary Material 5 (148.1KB, pdf)
Supplementary Material 6 (27KB, xlsx)
Supplementary Material 7 (17.1KB, xlsx)

Data Availability Statement

The relevant data can be found in the main text and supplementary materials.

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