Background/Objectives: Rice production faces threats from rising temperatures, demanding thermotolerant varieties. This study characterizes transcriptomic dynamics and identifies Hsp101-1 (heat shock protein 101-1)-associated gene regulatory modules in rice under reproductive-stage heat stress. Methods: Transcriptomics and WGCNA (weighted gene co-expression network analysis) were conducted in flag leaves and spikelets for wild-type (WT) and Hsp101-1-overexpressing (OE) lines under 40 °C stress at six time points (0-24 h) to reveal the change in gene expressions. Results: The number of DEGs (differentially expressed genes) revealed substantial pre-existing differences in WT and OE lines. Pre treatment, OE flag leaves showed 545 upregulated and 676 downregulated DEGs versus WT leaves. Post heat shock, the number of DEGs in flag leaves and spikelets was significantly reduced by 70-80%. KEGG enrichment of common DEGs across time points showed both WT and OE flag leaves enriched for ribosome biogenesis, ribosomes, and chaperones/folding catalysts. WGCNA identified that the MEdarkslateblue module correlated negatively with WT and positively with OE flag leaves. The MEturquoise module was suppressed at 1 h but activated by 8 h. Spikelet analysis identified the MElightpink4 module (negative correlation with WT, positive with OE) and a similarly dynamic MEturquoise module. Venn analysis identified 76 shared module genes, 71 of which were upregulated in the OE line, indicating that Hsp101-1 activates common protective targets. Hsp101-1's expression in the WT line was low basally, significantly upregulated at 1-8 h post shock, and returned to low levels by 24 h. Conclusions:Hsp101-1 enhances thermotolerance by (1) constitutively pre-stabilizing transcriptomic networks and reducing transcriptional fluctuations under heat stress and (2) modularly coordinating tissue-specific responses, providing a climate resilience framework.
Panicle architecture is an agronomic determinant of crop yield and a target for cereal crop improvement. To investigate its molecular mechanisms in rice, we performed map-based cloning and characterization of OPEN PANICLE 1 (OP1), a gain-of-function allele of LIGULELESS 1 (LG1), controlling the spread-panicle phenotype. This allele results from a 48-bp deletion in the LG1 upstream region and promotes pulvinus development at the base of the primary branch. Increased OP1 expression and altered panicle phenotype in chimeric transgenic plants and upstream-region knockout mutants indicated that the deletion regulates spread-panicle architecture in the mutant spread panicle 1 (sp1). Knocking out BRASSINOSTEROID UPREGULATED1 (BU1) gene in the background of OP1 complementary plants resulted in compact panicles, suggesting OP1 may regulate inflorescence architecture via the brassinosteroid signaling pathway. We regard that manipulating the upstream regulatory region of OP1 or genes involved in BR signal pathway could be an efficient way to improve rice inflorescence architecture.
With the increasing frequency of climatic anomalies, high temperatures and long-term rain often occur during the rice-harvesting period, especially for early rice crops in tropical and subtropical regions. Seed dormancy directly affects the resistance to pre-harvest sprouting (PHS). Therefore, in order to increase rice production, it is critical to enhance seed dormancy and avoid yield losses to PHS. The elucidation and utilization of the seed dormancy regulation mechanism is of great significance to rice production. Preliminary results indicated that the OsMKKK62-OsMKK3-OsMPK7/14 module might regulate ABA sensitivity and then control seed dormancy. The detailed mechanism is still unclear. The overexpression of OsMKK3 resulted in serious PHS. The expression levels of OsMKK3 and OsMPK7 were upregulated by ABA and GA at germination stage. OsMKK3 and OsMPK7 are both located in the nucleus and cytoplasm. The dormancy level of double knockout mutant mkk3/mft2 was lower than that of mkk3, indicating that OsMFT2 functions in the downstream of MKK3 cascade in regulating rice seeds germination. Biochemical results showed that OsMPK7 interacted with multiple core ABA signaling components according to yeast two-hybrid screening and luciferase complementation experiments, suggesting that MKK3 cascade regulates ABA signaling by modulating the core ABA signaling components. Moreover, the ABA response and ABA responsive genes of mpk7/14 were significantly higher than those of wild-type ZH11 when subjected to ABA treatment. MKK3 cascade mediates the negative feedback loop of ABA signal through the interaction between OsMPK7 and core ABA signaling components in rice.
Despite the crucial role of UDP-glycosyltransferases (UGTs) in various stress responses in plants, their biological functions in rice (Oryza sativa L.) remain poorly understood. In this study, we introduce a novel gene, OsUGT706E2, which is expressed at significantly higher levels in wild rice compared to cultivated rice. OsUGT706E2 is active in multiple tissues and localizes to both the nucleus and cytoplasm. Its transcription is notably up-regulated in response to cold stress, blast disease, and several hormone treatments. Overexpression of OsUGT706E2 markedly reduces seedling tolerance to blast disease, cold, and osmotic stress, whereas knocking out OsUGT706E2 enhances seedling tolerance to blast disease and osmotic stress. The expression levels of stress-related genes in OsUGT706E2 overexpressing plants were significantly lower compared to wild-type plants following stress treatment. Conversely, OsUGT706E2 knockout plants exhibited markedly higher expression levels of these genes than the control plants after stress treatment. Metabolome analysis further indicated that OsUGT760E2 influences metabolite content in rice. Specifically, OsUGT760E2-overexpressing seedlings had higher amino acid content and lower lipid content compared to wild-type seedlings, while OsUGT760E2-knockout seedlings showed lower amino acid content and higher lipid content than control seedlings. These findings suggest that OsUGT760E2 negatively regulates resistance to blast disease as well as tolerance to cold and osmotic stress in rice. As a result, OsUGT760E2 represents a promising target for enhancing rice stress tolerance through gene editing approaches.
The root-associated microbiota has a close relation to the life activities of plants, and its composition is affected by the rhizospheric environment and plant genotypes. Rice (Oryza sativa) was domesticated from the ancestor species Oryza rufipogon. Many important agricultural traits and adversity resistance of rice have changed during a long time of natural domestication and artificial selection. However, the influence of rice genotypes on root microbiota in important agricultural traits remains to be explained. In this study, we performed 16S rRNA and internal transcribed spacer (ITS) gene amplicon sequencing to generate bacterial and fungal community profiles of O. rufipogon and O. sativa, both of which were planted in a farm in Guangzhou and had reached the reproductive stage. We compared their root microbiota in detail by alpha diversity, beta diversity, different species, core microbiota, and correlation analyses. We found that the relative abundance of bacteria was significantly higher in the cultivated rice than in the common wild rice, while the relative abundance of fungi was the opposite. Significant differences in agricultural traits between O. rufipogon and O. sativa showed a high correlation with core microorganisms in the two Oryza species, which only existed in either or had obviously different abundance in both two species, indicating that rice genotype/phenotype had a strong influence on recruiting specific microorganisms. Our study provides a theoretical basis for the in-depth understanding of rice root microbiota and the improvement of rice breeding from the perspective of the interaction between root microorganisms and plants.IMPORTANCEPlant root microorganisms play a vital role not only in plant growth and development but also in responding the biotic and abiotic stresses. Oryza sativa is domesticated from Oryza rufipogon which has many excellent agricultural traits especially containing resistance to biotic and abiotic stresses. To improve the yield and resistance of cultivated rice, it is particularly important to deeply research on differences between O. sativa and O. rufipogon and find beneficial microorganisms to remodel the root microbiome of O. sativa.
Carbohydrates, proteins, lipids, minerals and vitamins are nutrient substances commonly seen in rice grains, but anthocyanidin, with benefit for plant growth and animal health, exists mainly in the common wild rice but hardly in the cultivated rice. To screen the rice germplasm with high intensity of anthocyanidins and identify the variations, we used metabolomics technique and detected significant different accumulation of anthocyanidins in common wild rice (Oryza rufipogon, with purple leaf sheath) and cultivated rice (Oryza sativa, with green leaf sheath). In this study, we identified and characterized a well-known MYB transcription factor, OsC1, through phenotypic (leaf sheath color) and metabolic (metabolite profiling) genome-wide association studies (pGWAS and mGWAS) in 160 common wild rice (O. rufipogon) and 151 cultivated (O. sativa) rice varieties. Transgenic experiments demonstrated that biosynthesis and accumulation of cyanidin-3-Galc, cyanidin 3-O-rutinoside and cyanidin O-syringic acid, as well as purple pigmentation in leaf sheath were regulated by OsC1. A total of 25 sequence variations of OsC1 constructed 16 functional haplotypes (higher accumulation of the three anthocyanidin types within purple leaf sheath) and 9 non-functional haplotypes (less accumulation of anthocyanidins within green leaf sheath). Three haplotypes of OsC1 were newly identified in our germplasm, which have potential values in functional genomics and molecular breeding of rice. Gene-to-metabolite analysis by mGWAS and pGWAS provides a useful and efficient tool for functional gene identification and omics-based crop genetic improvement.
In a genome-wide association study, we identified a rice UDP-glycosyltransferase gene, OsUGT706D2, whose transcription was activated in response to cold and submergence stress and to exogenous abscisic acid (ABA). OsUGT706D2 positively regulated the biosynthesis of tricin-4′-O-(syringyl alcohol) ether-7-O-glucoside at both the transcriptional and metabolic levels. OsUGT706D2 mediated cold and submergence tolerance by modulating the expression of stress-responsive genes as well as the abscisic acid (ABA) signaling pathway. Gain of function of OsUGT706D2 increased cold and submergence tolerance and loss of function of OsUGT706D2 reduced cold tolerance. ABA positively regulated OsUGT706D2-mediated cold tolerance but reduced submergence tolerance. These findings suggest the potential use of OsUGT706D2 for improving abiotic stress tolerance in rice.
Rice bacterial blight (BB), caused by Xanthomonas oryzae pv. oryzae (Xoo), is a major threat to rice production and food security. Exploring new resistance genes and developing varieties with broad-spectrum and high resistance has been a key focus in rice disease resistance research. In a preliminary study, rice cultivar Fan3, exhibiting high resistance to PXO99A and susceptibility to AH28, was developed by enhancing the resistance of Yuehesimiao (YHSM) to BB. This study performed a transcriptome analysis on the leaves of Fan3 and YHSM following inoculation with Xoo strains AH28 and PXO99A. The analysis revealed significant differential expression of 14,084 genes. Among the transcription factor (TF) families identified, bHLH, WRKY, and ERF were prominent, with notable differences in the expression of OsWRKY62, OsWRKY76, and OsbHLH6 across samples. Over 100 genes were directly linked to disease resistance, including nearly 30 NBS–LRR family genes. Additionally, 11 SWEET family protein genes, over 750 protein kinase genes, 63 peroxidase genes, and eight phenylalanine aminolysase genes were detected. Gene ontology (GO) analysis showed significant enrichment in pathways related to defense response to bacteria and oxidative stress response. Kyoto Encyclopedia of Genes and Genomes (KEGG) enrichment analysis indicated that differentially expressed genes (DEGs) were enriched in phenylpropanoid biosynthesis and diterpenoid biosynthesis pathways. Gene expression results from qRT-PCR were consistent with those from RNA-Seq, underscoring the reliability of the findings. Candidate genes identified in this study that may be resistant to BB, such as NBS–LRR family genes LOC_Os11g11960 and LOC_Os11g12350, SWEET family genes LOC_Os01g50460 and LOC_Os01g12130, and protein kinase-expressing genes LOC_Os01g66860 and LOC_Os02g57700, will provide a theoretical basis for further experiments. These results suggest that the immune response of rice to the two strains may be more concentrated in the early stage, and there are more up-regulated genes in the immune response of the high-resistant to PXO99A and medium-resistant to AH28, respectively, compared with the highly susceptible rice. This study offers a foundation for further research on resistance genes and the molecular mechanisms in Fan3 and YHSM.
Plant metabolites including anthocyanins play an important role in the growth of plants, as well as in regulating biotic and abiotic stress responses to the environment. Here we report comprehensive profiling of 3315 metabolites and a further metabolic-based genome-wide association study (mGWAS) based on 292,485 SNPs obtained from 311 rice accessions, including 160 wild and 151 cultivars. We identified hundreds of common variants affecting a large number of secondary metabolites with large effects at high throughput. Finally, we identified a novel gene namely OsLSC6 (Oryza sativa leaf sheath color 6), which encoded a UDP 3-O-glucosyltransferase and involved in the anthocyanin biosynthesis of Cyanidin-3-Galc (sd1825) responsible for leaf sheath color, and resulted in significant different accumulation of sd1825 between wild (purple) and cultivars (green). The results of knockout transgenic experiments showed that OsLSC6 regulated the biosynthesis and accumulation of sd1825, controlled the purple leaf sheath. Our further research revealed that OsLSC6 also confers resistance to cold stress during the seedling stage in rice. And we identified that a SNP in OsLSC6 was responsible for the leaf sheath color and chilling tolerance, supporting the importance of OsLSC6 in plant adaption. Our study could not only demonstrate that OsLSC6 is a vital regulator during anthocyanin biosynthesis and abiotic stress responses, but also provide a powerful complementary tool based on metabolites-to-genes analysis by mGWAS for functional gene identification andpromising candidate in future rice breeding and improvement.
Rice(Oryza sativa L.) is one of the most important food crops in the world, low temperature destabilizes the growth and development of rice, and causes lower yield or harvestless in worse cases, has becoming one of the major environmental stresses that decrease rice yield and quality. The use of agricultural protection technologies to reduce cold damage were not only time-consuming and laborious, but also ineffective.Therefore, the cultivation of rice varieties showing cold tolerance in practice is the optimal way to solve this problem. In breeding of cold-tolerant varieties, precise identification and evaluation of cold tolerance rice in conjugation with the exploitation and utilization of important cold-tolerant genes become of significance. In recent years,several cold tolerant QTLs or genes had been identified by map-based cloning, GWAS and QTL analysis,and their functional mechanisms have been investigated. This article reviews the cold resistance of rice, from the aspects of identification methods and periods, evaluation systems and the molecular research basis, and suggested that different evaluation systems should be used to evaluate the cold tolerance of rice at different growth stages. At the same time, it is proposed to pyramid important cold tolerance genes/QTLs in excellent rice germplasm resources for further innovative utilization, which could provide reference for exploring the mechanism of cold tolerance and breeding new varieties of cold tolerance in rice.
As the pioneer of the Green Revolution in China, Guangdong province witnessed the improvement and spread of semi-dwarf Xian/Indica rice cultivars and possessed diverse rice germplasm of landrace and cultivars. A total of 517 accessions containing a core germplasm of 479 newly sequenced landraces and modern cultivars were used to reveal breeding signatures and key variations for regional genetic improvement of indica rice from Guangdong. Four subpopulations were identified in the collection, which including Ind IV as a novel subpopulation that not covered by previously released accessions. Modern cultivars of subpopulation Ind II were inferred to have less deleterious variations, especially in yield related genes. About 15 Mb genomic segments were identified as potential breeding signatures by cross-population likelihood method (XP-CLR) of modern cultivars and landraces. The selected regions spanning multiple yield related QTLs (quantitative trait locus) which identified by GWAS (genome-wide association studies) of the same population, and specific variations that fixed in modern cultivars of Ind II were characterized. This study highlights genetic differences between traditional landraces and modern cultivars, which revealed the potential molecular basis of regional genetic improvement for Guangdong indica rice from southern China.
Rice blast caused by Magnaporthe oryzae is one of the most serious rice diseases worldwide. The early indica rice thermosensitive genic male sterile (TGMS) line HD9802S has the characteristics of stable fertility, reproducibility, a high outcrossing rate, excellent rice quality, and strong combining ability. However, this line exhibits poor blast resistance and is highly susceptible to leaf and neck blasts. In this study, backcross introduction, molecular marker-assisted selection, gene chipping, anther culture, and resistance identification in the field were used to introduce the broad-spectrum blast-resistance gene R6 into HD9802S to improve its rice blast resistance. Six induction media were prepared by varying the content of each component in the culture medium. Murashige and Skoog's medium with 3 mg/L 2,4-dichlorophenoxyacetic acid, 2 mg/L 1-naphthaleneacetic acid, and 1 mg/L kinetin and N6 medium with 800 mg/L casein hydrolysate, 600 mg/L proline, and 500 mg/L glutamine could improve the callus induction rate and have a higher green seedling rate and a lower white seedling rate. Compared to HD9802S, two doubled haploid lines containing R6 with stable fertility showed significantly enhanced resistance to rice blast and no significant difference in spikelet number per panicle, 1000-grain weight, or grain shape. Our findings highlight a rapid and effective method for improving rice blast resistance in TGMS lines.
This study investigates genetic variations of Ghd7 in natural rice populations. Our goal was to identify a solid theoretical basis for the flexible application of different Ghd7 alleles in rice genetic breeding. Genetic variations and haplotypes of Ghd7 were analyzed in 4326 cultivated varieties and eight accessions of common wild rice ( Oryza rufipogon) , and some of their heading date and plant height were investigated. The Ghd7 genome contains 79 single nucleotide polymorphisms (SNP) and one insertion/deletion length polymorphism (InDel), among which 10 SNPs caused non-synonymous mutations in amino acids. These were divided into 18 haplotypes, and there were two new non-synonymous mutations in the CCT domain of Ghd7 in O. rufipogon . In this study, a total of 10 new protein types of Ghd7 were found, and 3 new protein types were found only in O. rufipogon accessions. The cultivated rice was divided into indica and japonica subpopulations. There were significant differences in heading date, yield per plant, and plant height among the 8 haplotypes. The phenotype of heading date was significantly different only in Wuhan in 2018 between indica and japonica subpopulations. The Ghd7 allele of O. rufipogon S1111, which contains a higher plant height and a longest heading date, probably encodes a new strong functional protein type. There are abundant genetic variations of Ghd7 in natural populations, and it is possible to excavate new elite haplotypes of Ghd7 in O. rufipogon for breeding.
Floral organ development determines agricultural productivity by affecting seed development, seed quality, and final yield. In this study, we described the novel ogl mutant in rice (Oryza sativa L.), which is characterized by an open-glume phenotype, increased pistil number, reduced stamen number, decreased seed setting rate, and smaller rice grains. Genetic analysis showed that the open-glume phenotype might be controlled by a recessive qualitative trait locus. Employing bulked segregant analysis (BSA), one candidate region was identified on rice chromosome 1. The glume opening phenotype cosegregated with SNP (Chr1:1522703), which was located at the start codon of one transcript of OsJAG, resulting in partial loss of OsJAG function. cDNA analysis revealed that OsJAG encodes two transcript variants. Compared to normal plants, the expression of OsJAG.1 was upregulated in open-glume plants. When investigating the glume phenotype, we found that the expression of genes related to floral development changed greatly in open-glume plants. Taken together, this work increases our understanding of the developmental role of OsJAG in rice floral development.
Gamma-amino butyric acid (GABA) is a natural non-protein amino acid involved in stress, signal transmission, carbon and nitrogen balance, and other physiological processes in plants. In the human body, GABA has the effects of lowering blood pressure, anti-aging, and activating the liver and kidneys. However, there are few studies on the molecular regulation mechanism of genes in the metabolic pathways of GABA during grain development of giant embryo rice with high GABA content. In this study, three glant embryo (ge) mutants of different embryo sizes were obtained by CRISPR/Cas9 knockout, and it was found that GABA, protein, crude fat, and various mineral contents of the ge mutants were significantly increased. RNA-seq and qRT-PCR analysis showed that in the GABA shunt and polyamine degradation pathways, the expression levels of most of the genes encoding enzymes promoting GABA accumulation were significantly upregulated in the ge-1 mutant, whereas, the expression levels of most of the genes encoding enzymes involved GABA degradation were significantly downregulated in the ge-1 mutant. This is most likely responsible for the significant increase in GABA content of the ge mutant. These results help reveal the molecular regulatory network of GABA metabolism in giant embryo rice and provide a theoretical basis for the study of its development mechanisms, which is conducive to the rapid cultivation of GABA-rich rice varieties, promoting human nutrition, and ensuring health.
[Objective]It aims to study the yield performance and source-sink-flow characteristics of new strain of super-large-panicle rice DS23 in early season,and explore the source-flow-sink characteristics and super-high yield potential of DS23,with an aim to provide theoretical basis for high-yield breeding and cultivation of early rice in South China.[Method]The study was conducted under field conditions with the new super-large panicle type strain DS23 as test material and the medium to large panicle type variety YHSM as the control.The leaf area index,SPAD value,accumulation and transport characteristics of non-structural carbohydrate(NSC)in stems,anatomical characteristics of vascular bundles and yield component factors of different panicle types of rice were measured and compared.[Result]The number of grain per panicle,grain weight,sink capacity and yield of DS23 were increased by 37.3%,4.0%,33.7%and 15.9%respectively compared to those of YHSM seedlings.The effective number of panicles and seed setting rate were significantly decreased by 6.8%and 15.1%compared to those of YHSM seedlings.There was no significant difference in leaf area index and flag leaf SPAD value between the two varieties at heading stage.Compared with YHSM,total mass of NSC reserved in stem and apparent contribution of transferred NSC to grain yield of DS23 were significantly reduced by 15.3%and 19.8%,respectively,and the apparent transport rate of stem sheath NSC was significantly increased by 10.3%;The DS23 vascular bundle number was significantly increased by 17.3%,and there was no significant difference in the vascular bundle area between the two varieties;DS23 showed no significant change in grain/leaf ratio,with a significant decrease in NSC/spikelet ratio of 34.2%and an increase in vascular bundle load index of 20.4%-6 0.2%.[Conclusion]The super-large panicle type rice germplasm DS23 planted in early season has a large sink capacity,well-developed vascular bundles transport organization,strong material transformation ability in later stage and great potential for high yield and yield increase.Improvement in seed setting rate is a main direction for exploring its super-high yield potential.The study result lays a theoretical foundation for the breeding of super-large panicle early rice varieties and the research and development of high yield cultivation techniques.
萌发期耐冷性的强弱直接影响水稻直播时的成苗率.为高效地培育萌发期耐冷性强的水稻品种,本研究通过统计309份广东省水稻核心种质资源的低温萌发比率,筛选到57份萌发期耐冷性强的水稻品种.同时,以低温萌发比率为表型数据和高密度的SNP标记为基因型数据,利用全基因组关联分析鉴定到11个水稻低温萌发QTLs.其中qLTG2-1、qLTG7-1、qLTG7-2和qLTG10-1与前人定位的水稻低温萌发QTL共定位.qLTG11-1是本研究新鉴定的最显著的水稻低温萌发QTL.利用RNA-seq和qRT-PCR对其进行候选基因预测和验证,发现在低温萌发期间,qLTG11-1 区间内的LOC_Os11g07020在低温萌发比率高和低的品种中有不同的表达模式.LOC_Os11g07020编码果糖-1,6-二磷酸醛羧酶同工酶,可能通过调控糖酵解反应而影响水稻萌发期耐冷性.本研究筛选到多份耐冷性强的水稻品种,并找到可能调控水稻萌发期耐冷性的候选基因,可为培育萌发期耐冷性强的水稻品种提供参考.
自水稻矮化育种成功以来,广东的水稻品种在国内的水稻育种中发挥了重要作用.国家"十四五"规划明确提出了我国种业自主创新将以提升核心种源竞争力为目标,着力加强种质资源保护利用.本研究利用开发的10,610个多态性SLAF标签,对98份广东省不同时期育成的代表性品种和地方稻种,开展遗传演化、群体遗传结构及主成分分析,发现43份老品种和5份地方稻种的遗传基础较宽,而50份近期育成的新品种的遗传基础较窄;遗传演化和系谱分析发现,老品种大多数含有广场13、南特号、矮仔占、塘埔矮和竹印2号的血缘,而新品种大多含有IR系列品种、粳籼677、广场矮、中籼3588的血缘.这表明自20世纪90年代以来重视了外引材料的利用,但忽视了对本地优良种质资源的利用.这可能与20世纪70年代三系杂交稻的兴起有关.因此,为了确立核心种源的竞争地位,应重点加强本地优良种质资源的鉴评和利用.
穗发芽对籼稻生产危害严重.编辑休眠调控基因,创制高休眠性新种质,进而提高穗发芽抗性,是探索改良穗发芽抗性的新途径.泰丰B(TB)是籼型优质杂交稻保持系,主要缺点是种子休眠性偏低,易受穗发芽危害.MPK7/14具有抑制粳稻种子休眠作用,尚不清楚是否适用于籼稻品种.本研究利用CRISPR/Cas9技术编辑泰丰B(TB)的MPK7和MPK14基因,通过测序筛选获得纯合变异株系,并利用发芽试验分析变异种子的休眠特性.共获得6个转基因株系,从其后代中筛到2个纯合移码突变株系.纯合变异后代种子休眠性大幅提高,杂合变异后代的休眠性也有明显改善.与泰丰B(TB)相比,变异纯合后代株高、分蘖数、穗粒数差异不显著.综上所述,同时敲除MPK7和MPK14可以提高泰丰B(TB)种子休眠性,抑制穗发芽和解决水稻穗发芽问题.这些结果为进一步解析穗发芽的抗性机制以及探索新的育种改良方法提供了参考.