Abstract
As one of the most important staple foods, wheat plays a crucial role in sustaining food security. However, its production is greatly threatened by abiotic stresses such as salt (Khataar et al., 2018; Qi et al., 2019). Therefore, discovering the genes involved in abiotic stress tolerance and cultivating genetically modified plant varieties with enhanced stress tolerance are currently among the most important goals for plant breeders (Genc et al., 2019). To identify wheat transcriptional factor (TF) genes that are essential for regulating abiotic stress tolerance, 1455 wheat TF-encoding genes from full-length cDNA sequence libraries were tested for their expression under abiotic stress treatment conditions. One gene, TabZIP15 (TraesCS7A02G488600), was identified as a salt stress-responsive gene in wheat. TabZIP15 contains six exons and five introns in its structure (Figure 1a) and encodes a bZIP TF (Figure 1b). In the roots at the seedling stage, TabZIP15 expression was induced quickly and peaked within 1 h following salt stress treatment, and the expression level was approximately 2.4-fold greater than that before treatment (Figure 1c). TabZIP15 is a nuclear-localized protein (Figure 1d) with transcriptional activation activity in yeast (Figure 1e). Moreover, we also found that TabZIP15 has an affinity for the ABA-responsive element (ABRE) Cis-element using the yeast one-hybrid assay system (Figure 1f). To determine the role of the TabZIP15 gene in the regulation of salt stress tolerance in wheat, transgenic lines of the wheat variety Kenong 199 (KN199) overexpressing TabZIP15 under the control of the maize Ubi promoter were generated. Under normal conditions, no obvious phenotypic variation was detected between KN199 plants and the transgenic lines. However, after 20 days of salt stress treatment with 300 mm NaCl, although both the KN199 and transgenic plants displayed a certain degree of wilting and inhibited growth, the wilting degree was obviously lower in the transgenic plants than in the KN199 plants (Figure 1g). In addition, the transgenic plants presented obvious increases in height and aboveground fresh weight, and significant decreases in malondialdehyde (MDA) and H2O2 contents (Figure 1h-k). Considering that the root system is closely related to plant abiotic stress tolerance, we also examined the morphology of the roots and found no obvious differences in root morphology, length or fresh weight between the KN199 plants and transgenic plants under normal growth conditions. However, under salt stress conditions, although the root growth of both the transgenic plants and KN199 plants was inhibited, the transgenic plants had a larger root system than the KN199 plants (Figure 1l), and the root length and fresh weight were greater in the transgenic lines than in the KN199 plants (Figure 1m and n). In the reproductive stage, the spikes of the transgenic plants were almost the same as those of the KN199 plants under normal growth conditions, while the spike length was longer and the number of grains per spike was significantly higher in the transgenic plants than in the KN199 plants (Figure 1o-q). Together, these results suggest that the TabZIP15 gene is involved in the regulation of wheat salt stress tolerance. To better understand how TabZIP15 is involved in the wheat stress response, the truncated TabZIP15 was used to screen the yeast library. The results showed that the enolase protein TaENO-b (KC342470.1) could physically interact with TabZIP15 (Figure 1r). We then conducted luciferase complementation imaging and bimolecular fluorescence complementation assays in Nicotiana benthamiana and confirmed the interaction between TabZIP15 and TaENO-b (Figure 1s and t). TaENO-b is an enolase protein that catalyses the reversible dehydration of 2-phospho-D-glycerate to phosphoenolpyruvate as part of the glycolytic and gluconeogenesis pathways. Together, these results indicated that TabZIP15 may interact with enolase TaENO-b to participate in the regulation of the glycolysis and gluconeogenesis pathways, thus improving salt stress tolerance in wheat. To further investigate the putative mechanisms of the TabZIP15 gene, we used RNA sequencing (RNA-seq) transcriptomic analysis to identify differentially expressed genes (DEGs) (|log2 fold change [FC]| > 1, false discovery rate [FDR] < 0.05) between the KN199 and H381 transgenic plants before (0 h) and after salt stress treatment (1 h) at the seedling stage (Audic and Claverie, 1997; Mariani et al., 2003). As a result, a total of 3332 DEGs were found between KN199 and H381 plants at 0 h, including 1297 up-regulated and 2035 down-regulated DEGs in H381. At 1 h, 2807 DEGs were identified between KN199 and H381 plants, including 2691 up-regulated and 116 down-regulated DEGs in H381 plants (Figure 1u). We then tested the expression of four genes by qRT-PCR, and the results were consistent with the transcriptome analysis results (Figure 1v). Considering that the genes whose expression was specifically induced or inhibited after salt stress treatment may have more important roles in regulating salt tolerance in the transgenic plants, we further compared the DEGs between 0 h and 1 h. The results showed that the expression of 2556 DEGs was specifically up-regulated after salt treatment, while the expression of 45 DEGs was specifically down-regulated (Figure 1w). The large difference in the numbers of up-regulated and down-regulated genes may reflect TabZIP15’s role as a transcriptional activator. Gene Ontology (GO) enrichment analysis was performed for the DEGs that were specifically up-regulated at 1 h in transgenic plants. The 2556 genes were mainly enriched in 20 biological processes, including metabolic processes (GO:0008152), iron ion binding (GO:0005506) and the response to abiotic stimuli (GO:0009628; Figure 1x). Among these GO terms, the metabolic processes item enriched more than 900 DEGs, suggesting that the genes involved in metabolic regulation may play important roles in improving the salt tolerance of wheat through TabZIP15. Furthermore, more than 30 DEGs were enriched in the response to abiotic stimulus GO term. These DEGs may also regulate salt stress tolerance in wheat. In the future, we will analyse these DEGs in depth, which will help us to further understand the mechanism of the TabZIP15 gene regulating salt tolerance in wheat. Discovering stress-responsive genes and studying their roles via genetic modification approaches is necessary for the development of plants with enhanced stress tolerance (Abhinandan et al., 2018; Hasegawa et al., 2000; Zhu, 2016). In this study, we characterized a salt-induced bZIP gene, TabZIP15, in wheat. Based on the performance of TabZIP15 in transgenic plants, we propose that manipulating the expression of this gene leads to enhanced salt stress tolerance. These results improve our understanding of the roles of wheat bZIP protein in the plant response to abiotic stress and highlight a candidate gene for wheat improvement. We thank Jizeng Jia and Ruilian Jing's groups for providing FLC and yeast library. This work was supported by the National Science Foundation (31771792), the National Transgenic Research Project (2016ZX08009001-001-004), the Agricultural Science and Technology Innovation Program of Chinese Academy of Agricultural Sciences and the Tianshan Innovation Team Plan (2020D14002). The authors declare no conflict of interest. X.K. and L.Z. conceived the study and designed the experiments. C.B., C.D., Y.Y., C.J., Y.Y., Y.Z., C.X. and F.D. performed the experiments. L.Z., X.K., Z.N., C.B., Y.Y. and Y.Y. wrote the manuscript.
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Claims 20
Claim · model-written summary
TabZIP15 was identified as salt responsive in wheat.
- Taxon
- wheat
Triticum aestivume0 - Gene
- TabZIP15e0
- Outcome
- salt stress-responsive genee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“One gene, TabZIP15 (TraesCS7A02G488600), was identified as a salt stress-responsive gene in wheat.”
Claim · model-written summary
TabZIP15 has six exons and five introns and encodes a bZIP transcription factor.
- Taxon
- wheatTriticum aestivumt0
- Gene
- TabZIP15e0
- Outcome
- six exons and five intronse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“TabZIP15 contains six exons and five introns in its structure (Figure 1a) and encodes a bZIP TF”
t0“wheat”
Claim · model-written summary
Seedling root TabZIP15 expression peaked within an hour of salt treatment at about 2.4 times pretreatment expression.
- Taxon
- wheatTriticum aestivumt0
- Organ / stage
- roots at the seedling stagee0
- Treatment
- salt stress treatmente0
- Gene
- TabZIP15e0
- Outcome
- TabZIP15 expressione0
- Direction
- greatere0
- Value
- 2.4-folde0
- Comparator
- before treatmente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“In the roots at the seedling stage, TabZIP15 expression was induced quickly and peaked within 1 h following salt stress treatment, and the expression level was approximately 2.4-fold greater than that before treatment”
t0“wheat”
Claim · model-written summary
TabZIP15 is nuclear localized and activates transcription in yeast.
- Taxon
- wheatTriticum aestivumt0
- Gene
- TabZIP15e0
- Outcome
- nuclear-localized proteine0
- Qualifier
- transcriptional activation activity in yeaste0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“TabZIP15 is a nuclear-localized protein (Figure 1d) with transcriptional activation activity in yeast”
t0“wheat”
Claim · model-written summary
TabZIP15 binds the ABA-responsive element in a yeast assay.
- Taxon
- wheatTriticum aestivumt0
- Gene
- TabZIP15e0
- Outcome
- affinity for the ABA-responsive element (ABRE) Cis-elemente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“TabZIP15 has an affinity for the ABA-responsive element (ABRE) Cis-element using the yeast one-hybrid assay system”
t0“wheat”
Claim · model-written summary
Under normal conditions, transgenic and KN199 wheat had no obvious phenotypic difference.
- Taxon
- wheatTriticum aestivumt0
- Treatment
- transgenic linese0
- Outcome
- phenotypic variatione0
- Direction
- no obviouse0
- Qualifier
- Under normal conditionse0
- Comparator
- KN199 plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“Under normal conditions, no obvious phenotypic variation was detected between KN199 plants and the transgenic lines.”
t0“wheat”
Claim · model-written summary
After 20 days of 300 mm NaCl, TabZIP15 transgenic wheat wilted less than KN199.
- Taxon
- wheatTriticum aestivumt0
- Treatment
- transgenic plantse0
- Outcome
- wilting degreee0
- Direction
- lowere0
- Qualifier
- after 20 days of salt stress treatment with 300 mm NaCle0
- Comparator
- KN199 plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“after 20 days of salt stress treatment with 300 mm NaCl, although both the KN199 and transgenic plants displayed a certain degree of wilting and inhibited growth, the wilting degree was obviously lower in the transgenic plants than in the KN199 plants”
t0“wheat”
Claim · model-written summary
Salt treated transgenic wheat had greater height and aboveground fresh weight.
- Taxon
- wheatTriticum aestivumt0
- Treatment
- transgenic plantse0
- Outcome
- height and aboveground fresh weighte0
- Direction
- increasese0
Not reportedComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“the transgenic plants presented obvious increases in height and aboveground fresh weight”
t0“wheat”
Claim · model-written summary
Salt treated transgenic wheat had lower malondialdehyde and H2O2 contents.
- Taxon
- wheatTriticum aestivumt0
- Outcome
- malondialdehyde (MDA) and H2O2 contentse0
- Direction
- decreasese0
Not reportedTreatmentComparator
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“significant decreases in malondialdehyde (MDA) and H2O2 contents”
t0“wheat”
Claim · model-written summary
Under normal conditions, root morphology, length and fresh weight did not obviously differ between transgenic and KN199 wheat.
- Taxon
- wheatTriticum aestivumt0
- Treatment
- transgenic plantse0
- Outcome
- root morphology, length or fresh weighte0
- Direction
- no obvious differencese0
- Qualifier
- normal growth conditionse0
- Comparator
- KN199 plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“no obvious differences in root morphology, length or fresh weight between the KN199 plants and transgenic plants under normal growth conditions”
t0“wheat”
Claim · model-written summary
Under salt stress, transgenic wheat had a larger root system and greater root length and fresh weight than KN199.
- Taxon
- wheatTriticum aestivumt0
- Treatment
- transgenic plantse0
- Outcome
- root length and fresh weighte0
- Direction
- greatere0
- Comparator
- KN199 plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“the transgenic plants had a larger root system than the KN199 plants (Figure 1l), and the root length and fresh weight were greater in the transgenic lines than in the KN199 plants”
t0“wheat”
Claim · model-written summary
Transgenic and KN199 wheat spikes were almost the same under normal growth.
- Taxon
- wheatTriticum aestivumt0
- Treatment
- transgenic plantse0
- Outcome
- spikese0
- Direction
- almost the samee0
- Qualifier
- normal growth conditionse0
- Comparator
- KN199 plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“the spikes of the transgenic plants were almost the same as those of the KN199 plants under normal growth conditions”
t0“wheat”
Claim · model-written summary
Transgenic wheat had longer spikes and significantly more grains per spike than KN199.
- Taxon
- wheatTriticum aestivumt0
- Treatment
- transgenic plantse0
- Outcome
- the number of grains per spikee0
- Direction
- highere0
- Comparator
- KN199 plantse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“the spike length was longer and the number of grains per spike was significantly higher in the transgenic plants than in the KN199 plants”
t0“wheat”
Claim · model-written summary
TaENO-b physically interacted with TabZIP15, including confirmation in Nicotiana benthamiana.
- Taxon
- wheatTriticum aestivumt0
- Gene
- TabZIP15e0
- Outcome
- affinity for the ABA-responsive elemente0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“we also found that TabZIP15 has an affinity for the ABA-responsive element”
t0“wheat”
Claim · model-written summary
At zero hours, H381 and KN199 differed in 3332 genes, with 1297 upregulated and 2035 downregulated in H381.
- Taxon
- wheatTriticum aestivumt0
- Material
- H381e0
- Outcome
- DEGse0
- Value
- 3332 DEGse0
- Qualifier
- at 0 he0
- Comparator
- KN199e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“a total of 3332 DEGs were found between KN199 and H381 plants at 0 h, including 1297 up-regulated and 2035 down-regulated DEGs in H381”
t0“wheat”
Claim · model-written summary
At one hour, H381 and KN199 differed in 2807 genes, with 2691 upregulated and 116 downregulated in H381.
- Taxon
- wheatTriticum aestivumt0
- Material
- H381e0
- Outcome
- DEGse0
- Value
- 2807 DEGse0
- Qualifier
- At 1 he0
- Comparator
- KN199e0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“At 1 h, 2807 DEGs were identified between KN199 and H381 plants, including 2691 up-regulated and 116 down-regulated DEGs in H381 plants”
t0“wheat”
Claim · model-written summary
After salt treatment, 2556 differentially expressed genes were specifically upregulated and 45 downregulated.
- Taxon
- wheatTriticum aestivumt0
- Treatment
- salt treatmente0
- Outcome
- expression of 2556 DEGse0
- Direction
- up-regulatede0
- Value
- 2556 DEGse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“the expression of 2556 DEGs was specifically up-regulated after salt treatment, while the expression of 45 DEGs was specifically down-regulated”
t0“wheat”
Claim · model-written summary
The 2556 upregulated genes were enriched in 20 biological processes, including metabolism, iron ion binding and abiotic response.
- Taxon
- wheatTriticum aestivumt0
- Outcome
- enriched in 20 biological processese0
- Value
- 20 biological processese0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“The 2556 genes were mainly enriched in 20 biological processes, including metabolic processes (GO:0008152), iron ion binding (GO:0005506) and the response to abiotic stimuli (GO:0009628”
t0“wheat”
Claim · model-written summary
More than 900 genes were enriched in metabolic processes and more than 30 in abiotic stimulus response.
- Taxon
- wheatTriticum aestivumt0
- Outcome
- metabolic processes iteme0
- Value
- more than 900 DEGse0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“the metabolic processes item enriched more than 900 DEGs”
t0“wheat”
Claim · model-written summary
The authors propose that manipulating TabZIP15 expression enhances wheat salt tolerance.
- Taxon
- wheatTriticum aestivumt0
- Outcome
- manipulating the expression of this gene leads to enhanced salt stress tolerancee0
Organisms pictured · Representative pictures of the matched species, not figures from the paper.
Verbatim
e0“we propose that manipulating the expression of this gene leads to enhanced salt stress tolerance”
t0“wheat”
The same questions in other papers
salinity · molecular: expression · Triticum aestivum (wheat)Results differ
21 papers in the same direction as this paper · 17 in the opposite direction
- negative
- no effect
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