油菜角果热应答基因鉴定和拟南芥热激转录因子互作分析
|
摘要
高温是影响农业生产的非生物胁迫因子之一。长江流域是我国油菜的主产区,该地区在种子灌浆成熟期经常遭遇短期或连续30度以上的高温,造成种子高温逼熟,严重影响种子产量和含油量。但目前还缺乏对油菜耐高温逼熟的遗传和分子基础的了解。模式植物拟南芥中热激转录因子研究已取得重要进展,但如何有效应用到作物遗传改良还有待探索。针对这些问题,本研究主要开展了三个方面的工作：1)以开花20d后的油菜果皮和种子为材料,利用油菜95K表达序列芯片分析两种组织在热处理后24h和48h基因表达变化的情况,为研究油菜抗热反应提供更广泛的基因源。2)利用拟南芥热激转录因子多突变体,从表型、表达调控和蛋白互作分析以及遗传调控途径等方面研究HsfA与HsfB两个亚家族基因的功能和潜在的调控关系。3)根据油菜热激表达谱结果,以受热胁迫诱导表达的拟南芥基因H38(AT4G23493)为分析对象,从基因表达、亚细胞定位、生理生化反应以及生长发育调节等方面研究了其功能。主要研究结论如下：
1.在热处理后的油菜芯片中共检测到1248个上调和898个下调表达的基因,其中在果皮中检测到925个上调和581个下调表达基因,在种子中共检测到837个上调和383个下调表达基因。在果皮和种子中,这些转录水平的变化可能为揭示油菜角果期抗热的分子机制提供线索。
2.上调基因中有40.9%(511)同时存在于种子和果皮中,然而下调表达的基因中只有66个同时存在于两个组织中,并且上调基因的平均变化倍数(2.1-72.8)比下调(2.0-5.6)的更加剧烈。在共同诱导表达的基因中,与胁迫相关的基因以及与RNA和蛋白质相关等基因显著富集,其中包括热胁迫反应的标志性基因如Hsfs、 Hsps、DREB2a、ROF2、GolSl和MBF1c等,以及CYP707A4和细胞色素bd泛醌氧化酶等以前未鉴定到的基因。
3.在果皮中共检测到411个上调和514个下调表达基因,而这些基因在种子中的表达没有受到影响。其中,最突出的是21个参与硫苷代谢的基因被同时下调表达,另外一些表达改变的基因则参与光合作用,水、糖分以及离子转运等几个主要代谢途径,这些基因表达量的变化可能与热处理后油菜种子产量以及品质直接相关。
4.在种子中检测到325个上调和314个下调表达基因,其在果皮中的表达没有明显变化。与果皮类似,我们共检测到12个花青素合成途径上的基因表达受到抑制,此外与种子储藏蛋白、植物激素、油脂代谢等重要代谢途径相关的基因的表达也发生变化。
5.在所有的差异表达基因中,有1/3编码功能未知蛋白(果皮中有484个,种子中有398个),其中502个在拟南芥中能找到同源序列,181个则为油菜特有基因。因此,在植物中可能还存在许多与抗热相关的功能未知基因以及物种特有基因。
6.根据芯片数据,我们挑选了8个基因并获得了相应的拟南芥突变体。在种子萌发、幼苗、成熟植株(开花之前)等多个时期对这些突变体进行热处理,发现h25、h29与h38至少在一种热处理条件下表现热敏感。
7.通过对双突变体的分析发现,HsfA3位于HsfA2下游协同调节Hsp18.1和Hsp25.3-P等的表达。采用酵母双杂交的方法检测到HsfA2与HsfA3可以通过各自的OD结构域进行互作。利用HsfB的双突变体和三突变体,从抗热表型和靶标基因的表达变化等角度阐述了HsfB的功能及其之间的关系。
8.在芯片中发现的热应答基因H38(AT4G23493),在植物界中保守的。在拟南芥中,H38主要在成熟的种子以及萌发过程中高丰度表达；其编码的蛋白定位于线粒体。启动子和基因表达分析发现H38在幼苗以及发育的角果中能显著被热诱导。h38突变体在苗期对热处理敏感,而在抽薹开花期热处理时能够显著积累花青素。
9.采用生物信息学预测发现H38与GolS1、SK1、热激蛋白AT5G37670和AT2G19310等基因共表达,采用荧光定量的方法分析了共表达基因以及抗热的核心基因Hsf和Hsp和线粒体定位的抗热关键基因的表达变化,明确了H38的功能与调控网络。
10.h38突变体的种子在萌发过程中对葡萄糖和ABA的耐受性显著增强,这种耐受性与渗透胁迫无关。此外,h38突变体的花器官、种子等显著比野生型增大,细胞学观察表明器官增大主要是由于细胞体积增大造成。
High temperature stress is one of the major abiotic stresses in the world that severely restricts crop production. The Yangzi River Valley is the most important rapeseed oil production area in China, but this area usually suffers transitory or long period high temperature of over30℃at seed filling stage, which could result in heat-forced maturity, decreased oilseed yield and quality. However, the inheritance and molecular mechanisms of anti-heat-forced maturity in oilseed rape are still largely unknown. Heat resistance genes including heat stress transcription factors were well studied in model plant Arabidopsis, but how to apply the theoretical knowledge to genetically improve heat resistance of crops effectively is poorly understood. To address these issues, three major works were conducted in this study:1) Global transcription profiles in siliques of Brassica napus from20days after flowering after heat stress was analyzed using a95k Brassica60-mer GeneChip. The newly identified transcripts further enriched the reservoir of heat-responsive genes.2) Using double and triple mutants, functional analysis and potential regulation network of HsfAs and HsfBs were conducted in Arabidopsis by phenotypic analysis, gene expression and protein interaction.3) Functions of an unknown gene of H38(AT4G23493) that was discovered to be a potential heat regulator by microarray survey, were systematically studied from the aspects of gene expression, subcellular localization, physiological and biochemical studies and growth and developmental regulation analysis. The main conclusions of this study achieved are as follows:
1. We detected1248up-regulated and898down-regulated genes in the heat stressed rapeseed siliques;925up-regulated and581down-regulated were detected in the silique walls (SW), and837up-regulated and383down-regulated genes in the seeds. The alteration of the transcripts may provide clues for revealing the mechanism for heat-resistance in the oilseed rape at the seed filling stage.
2.40.9%(511) of the up-regulated genes are present in the seeds and silique walls simultaneously, however, only66genes were decreased in both organs; and the average fold-change of the up-regulated genes (2.1to72.8) was significantly larger than that of the down-regulated ones (2.0to5.6). Notably, stress-related genes and transcription factors were significantly enriched in both organs. Many of the up-regulated genes are known to be heat-marker genes, including13Hsfs and91Hsps, and DREB2a, ROF2, GolSl, MBFlc, and others such as CYP707A4and panthenol oxidized cytochrome bd enzyme which were not identified in heat response previously.
3.411up-regulated and514down-regulated genes were detected in the silique walls, but not affected in seeds. Among them, some of the genes were involved in the major metabolic pathways of photosynthesis and transport (water, sugar and ion) and a cluster of21genes that are involving in glucosinolates metabolism were simultaneously reduced in heat stressed silique walls. All these altered expression may directly related to the decreased yield and quality of rapeseed after heat stress.
4.325up-regulated and314down-regulated genes were altered in seeds and whose expression did not change significantly in the silique walls. Genes related to seed storage proteins, phytohormones, lip id metabolism and other important metabolic pathways were changed. Similar to the silique walls, we have detected as many as12genes that involved in the anthocyanin biosynthetic pathway were suppressed in heat stressed seeds.
5.1/3of all the differentially expressed genes encodes protein with unknown functions (484in silique walls,398in seeds),502of which can be found their homologous sequences in Arabidopsis, while181were rapeseed specific genes. Therefore, there may be many unknown and species-specific genes that regulate plant thermotolerance.
6. According to the microarray data,8genes were selected for further studies. Mutants of homozygous genes in Arabidopsis were obtained and confirmed by PCR and gene expression analysis. Further, thermotolerance assays were carried out at the stage of seed germination, young seedling and mature plants (before flowering) of each mutant; and h25, h29and h38were found to be heat-sensitive in at least one of the heat-treatment conditions.
7. From the phenotypic and expression analysis with the double mutants and single mutants of hsfA2and hsfA3under long-time recovery from heat stress showed that HsfA3was downstream of HsfA2and synergistically regulated the expression of HSP18.1and Hsp25.3-P. Moreover, HsfA2and HsfA3could strongly interact though their oligomerization domains by yeast two hybrid approach. Function analysis and regulatory study of three HsfBs were performed by phenotypic analysis after heat stress, and gene expression analysis of heat resistant symbol genes and potential target genes using double and triple mutants.
8. H38was identified as a conserved gene with unknown function in the plant kingdom. In Arabidopsis, H38was mainly expressed in the mature and germination seeds and encodes the protein that was located in mitochondrial. H38was significantly induced after heat stress in seedlings and developing siliques by histochemical staining and qRT-PCR analysis, Mutant of H38was sensitive to heat stress at seedlings stage; while it accumulates more anthocyanins after heat stress at the early stage of flowering.
9. Co-expression analysis revealed that expression of H38was intimately correlated to that of GolSl, SKI, heat shock protein AT5G37670and AT2G19310. The co-expression genes together with some representative Hsfs/Hsps and mitochondrial localized heat-related genes were studied by qRT-PCR, which provides important information of the regulatory network of H38.
10. The tolerance of seed germination under glucose or/and ABA treatment was significantly enhanced in h38mutant; what is more the floral organs and seeds of h38were significant larger than the wild type which was mainly resulted from the increased cell size by cytology observations.
1.在热处理后的油菜芯片中共检测到1248个上调和898个下调表达的基因,其中在果皮中检测到925个上调和581个下调表达基因,在种子中共检测到837个上调和383个下调表达基因。在果皮和种子中,这些转录水平的变化可能为揭示油菜角果期抗热的分子机制提供线索。
2.上调基因中有40.9%(511)同时存在于种子和果皮中,然而下调表达的基因中只有66个同时存在于两个组织中,并且上调基因的平均变化倍数(2.1-72.8)比下调(2.0-5.6)的更加剧烈。在共同诱导表达的基因中,与胁迫相关的基因以及与RNA和蛋白质相关等基因显著富集,其中包括热胁迫反应的标志性基因如Hsfs、 Hsps、DREB2a、ROF2、GolSl和MBF1c等,以及CYP707A4和细胞色素bd泛醌氧化酶等以前未鉴定到的基因。
3.在果皮中共检测到411个上调和514个下调表达基因,而这些基因在种子中的表达没有受到影响。其中,最突出的是21个参与硫苷代谢的基因被同时下调表达,另外一些表达改变的基因则参与光合作用,水、糖分以及离子转运等几个主要代谢途径,这些基因表达量的变化可能与热处理后油菜种子产量以及品质直接相关。
4.在种子中检测到325个上调和314个下调表达基因,其在果皮中的表达没有明显变化。与果皮类似,我们共检测到12个花青素合成途径上的基因表达受到抑制,此外与种子储藏蛋白、植物激素、油脂代谢等重要代谢途径相关的基因的表达也发生变化。
5.在所有的差异表达基因中,有1/3编码功能未知蛋白(果皮中有484个,种子中有398个),其中502个在拟南芥中能找到同源序列,181个则为油菜特有基因。因此,在植物中可能还存在许多与抗热相关的功能未知基因以及物种特有基因。
6.根据芯片数据,我们挑选了8个基因并获得了相应的拟南芥突变体。在种子萌发、幼苗、成熟植株(开花之前)等多个时期对这些突变体进行热处理,发现h25、h29与h38至少在一种热处理条件下表现热敏感。
7.通过对双突变体的分析发现,HsfA3位于HsfA2下游协同调节Hsp18.1和Hsp25.3-P等的表达。采用酵母双杂交的方法检测到HsfA2与HsfA3可以通过各自的OD结构域进行互作。利用HsfB的双突变体和三突变体,从抗热表型和靶标基因的表达变化等角度阐述了HsfB的功能及其之间的关系。
8.在芯片中发现的热应答基因H38(AT4G23493),在植物界中保守的。在拟南芥中,H38主要在成熟的种子以及萌发过程中高丰度表达；其编码的蛋白定位于线粒体。启动子和基因表达分析发现H38在幼苗以及发育的角果中能显著被热诱导。h38突变体在苗期对热处理敏感,而在抽薹开花期热处理时能够显著积累花青素。
9.采用生物信息学预测发现H38与GolS1、SK1、热激蛋白AT5G37670和AT2G19310等基因共表达,采用荧光定量的方法分析了共表达基因以及抗热的核心基因Hsf和Hsp和线粒体定位的抗热关键基因的表达变化,明确了H38的功能与调控网络。
10.h38突变体的种子在萌发过程中对葡萄糖和ABA的耐受性显著增强,这种耐受性与渗透胁迫无关。此外,h38突变体的花器官、种子等显著比野生型增大,细胞学观察表明器官增大主要是由于细胞体积增大造成。
High temperature stress is one of the major abiotic stresses in the world that severely restricts crop production. The Yangzi River Valley is the most important rapeseed oil production area in China, but this area usually suffers transitory or long period high temperature of over30℃at seed filling stage, which could result in heat-forced maturity, decreased oilseed yield and quality. However, the inheritance and molecular mechanisms of anti-heat-forced maturity in oilseed rape are still largely unknown. Heat resistance genes including heat stress transcription factors were well studied in model plant Arabidopsis, but how to apply the theoretical knowledge to genetically improve heat resistance of crops effectively is poorly understood. To address these issues, three major works were conducted in this study:1) Global transcription profiles in siliques of Brassica napus from20days after flowering after heat stress was analyzed using a95k Brassica60-mer GeneChip. The newly identified transcripts further enriched the reservoir of heat-responsive genes.2) Using double and triple mutants, functional analysis and potential regulation network of HsfAs and HsfBs were conducted in Arabidopsis by phenotypic analysis, gene expression and protein interaction.3) Functions of an unknown gene of H38(AT4G23493) that was discovered to be a potential heat regulator by microarray survey, were systematically studied from the aspects of gene expression, subcellular localization, physiological and biochemical studies and growth and developmental regulation analysis. The main conclusions of this study achieved are as follows:
1. We detected1248up-regulated and898down-regulated genes in the heat stressed rapeseed siliques;925up-regulated and581down-regulated were detected in the silique walls (SW), and837up-regulated and383down-regulated genes in the seeds. The alteration of the transcripts may provide clues for revealing the mechanism for heat-resistance in the oilseed rape at the seed filling stage.
2.40.9%(511) of the up-regulated genes are present in the seeds and silique walls simultaneously, however, only66genes were decreased in both organs; and the average fold-change of the up-regulated genes (2.1to72.8) was significantly larger than that of the down-regulated ones (2.0to5.6). Notably, stress-related genes and transcription factors were significantly enriched in both organs. Many of the up-regulated genes are known to be heat-marker genes, including13Hsfs and91Hsps, and DREB2a, ROF2, GolSl, MBFlc, and others such as CYP707A4and panthenol oxidized cytochrome bd enzyme which were not identified in heat response previously.
3.411up-regulated and514down-regulated genes were detected in the silique walls, but not affected in seeds. Among them, some of the genes were involved in the major metabolic pathways of photosynthesis and transport (water, sugar and ion) and a cluster of21genes that are involving in glucosinolates metabolism were simultaneously reduced in heat stressed silique walls. All these altered expression may directly related to the decreased yield and quality of rapeseed after heat stress.
4.325up-regulated and314down-regulated genes were altered in seeds and whose expression did not change significantly in the silique walls. Genes related to seed storage proteins, phytohormones, lip id metabolism and other important metabolic pathways were changed. Similar to the silique walls, we have detected as many as12genes that involved in the anthocyanin biosynthetic pathway were suppressed in heat stressed seeds.
5.1/3of all the differentially expressed genes encodes protein with unknown functions (484in silique walls,398in seeds),502of which can be found their homologous sequences in Arabidopsis, while181were rapeseed specific genes. Therefore, there may be many unknown and species-specific genes that regulate plant thermotolerance.
6. According to the microarray data,8genes were selected for further studies. Mutants of homozygous genes in Arabidopsis were obtained and confirmed by PCR and gene expression analysis. Further, thermotolerance assays were carried out at the stage of seed germination, young seedling and mature plants (before flowering) of each mutant; and h25, h29and h38were found to be heat-sensitive in at least one of the heat-treatment conditions.
7. From the phenotypic and expression analysis with the double mutants and single mutants of hsfA2and hsfA3under long-time recovery from heat stress showed that HsfA3was downstream of HsfA2and synergistically regulated the expression of HSP18.1and Hsp25.3-P. Moreover, HsfA2and HsfA3could strongly interact though their oligomerization domains by yeast two hybrid approach. Function analysis and regulatory study of three HsfBs were performed by phenotypic analysis after heat stress, and gene expression analysis of heat resistant symbol genes and potential target genes using double and triple mutants.
8. H38was identified as a conserved gene with unknown function in the plant kingdom. In Arabidopsis, H38was mainly expressed in the mature and germination seeds and encodes the protein that was located in mitochondrial. H38was significantly induced after heat stress in seedlings and developing siliques by histochemical staining and qRT-PCR analysis, Mutant of H38was sensitive to heat stress at seedlings stage; while it accumulates more anthocyanins after heat stress at the early stage of flowering.
9. Co-expression analysis revealed that expression of H38was intimately correlated to that of GolSl, SKI, heat shock protein AT5G37670and AT2G19310. The co-expression genes together with some representative Hsfs/Hsps and mitochondrial localized heat-related genes were studied by qRT-PCR, which provides important information of the regulatory network of H38.
10. The tolerance of seed germination under glucose or/and ABA treatment was significantly enhanced in h38mutant; what is more the floral organs and seeds of h38were significant larger than the wild type which was mainly resulted from the increased cell size by cytology observations.
引文
1.万兵喜.拟南芥T-DNA突变体对几种非生物胁迫处理的反应.[硕士学位论文].武汉华中农业大学图书馆：2008
2. 王汉中,华玮,刘静,刘贵华,王新发,杨庆.油菜热激蛋白基因HSP17.8及其应用.中国专利,101544983.2009-09-30
3. 张树杰,王汉中.我国油菜生产应对气候变化的对策和措施分析.中国油料作物学报,2012,34：114-122
4. Agarwal M, Katiyar-Agarwal S, Sahi C, Gallie DR, Grover A. Arabidopsis thaliana Hsp100 proteins:kith and kin. Cell Stress Chaperones,2001,6:219
5. Aksouh-Harradj N, Campbell L, Mailer R. Canola response to high and moderately high temperature stresses during seed maturation. Can J Plant Sci,2006,86: 967-980
6. Anderson LE. Hoyer's solution as a rapid permanent mounting medium for bryophytes. Bryologist,1954,57:242-244
7. Angadi S, Cutforth H, Miller P, McConkey B, Entz M, Brandt S, Volkmar K. Response of three Brassica species to high temperature stress during reproductive growth. Can J Plant Sci,2000,80:693-701
8. Baniwal SK, Chan KY, Scharf K-D, Nover L. Role of heat stress transcription factor HsfA5 as specific repressor of HsfA4. J Biol Chem,2007,282:3605-3613
9. Battisti DS, Naylor RL. Historical warnings of future food insecurity with unprecedented seasonal heat. Science,2009,323:240-244
10. Bharti K, von Ko skull-Dor ing P, Bharti S, Kumar P, Tintschl-Korbitzer A, Treuter E, Nover L. Tomato heat stress transcription factor HsfB1 represents a novel type of general transcription coactivator with a histone-like motif interacting with the plant CREB binding protein ortholog HAC1. Plant Cell,2004,16:1521-1535
11. Busch W, Wunderlich M, Schoffl F. Identification of novel heat shock factor-dependent genes and biochemical pathways in Arabidopsis thaliana. Plant J, 2004,41:1-14
12. Charng YY, Liu HC, Liu NY, Chi WT, Wang CN, Chang SH, Wang TT. A heat-inducible transcription factor, HsfA2, is required for extension of acquired thermotolerance in Arabidopsis. Plant Physiol,2007,143:251-262
13. Charng YY, Liu HC, Liu NY, Hsu FC, Ko SS. Arabidopsis Hsa32, a novel heat shock protein, is essential for acquired thermotolerance during long recovery after acclimation. Plant Physiol,2006,140:1297-1305
14. Chauhan H, Khurana N, Agarwal P, Khurana JP, Khurana P. A seed preferential heat shock transcription factor from wheat provides abiotic stress tolerance and yield enhancement in transgenic Arabidopsis under heat stress environment. PLoS ONE, 2013,8:e79577
15. Chauhan H, Khurana N, Tyagi AK, Khurana JP, Khurana P. Identification and characterization of high temperature stress responsive genes in bread wheat (Triticum aestivum L.) and their regulation at various stages of development. Plant Mol Biol,2011,75:35-51
16. Chen G, Bi YR, Li N. EGY1 encodes a membrane-associated and ATP-independent metalloprotease that is required for chloroplast development. Plant J,2005,41: 364-3751
17. Chen H, Hwang JE, Lim CJ, Kim DY, Lee SY, Lim CO. Arabidopsis DREB2C functions as a transcriptional activator of HsfA3 during the heat stress response. Biochem Biophys Res Commun,2010,401:238-244
18. Chen X, Ye C, Lii H, Xu M, Li W, Zhang L, Wang C, Luo S, Zhu B. Cloning of GmHSFAl gene and its overexpression leading to enhancement of heat tolerance in transgenic soybean. Yi Chuan,2006,28:1411-1420
19. Clarke SM, Cristescu SM, Miersch O, Harren FJM, Wasternack C, Mur LAJ. Jasmonates act with salicylic acid to confer basal thermotolerance in Arabidopsis thaliana. New Phytol, 2009,182:175-187
20. Clarke SM, Mur LAJ, Wood JE, Scott IM. Salicylic acid dependent signaling promotes basal thermotolerance but is not essential for acquired thermotolerance in Arabidopsis thaliana. Plant J,2004,38:432-447
21. Claros MG, Vincens P. Computational method to predict mitochondrially imported proteins and their targeting sequences. Eur J Biochem, 1996,241:779-786
22. Clough SJ, Bent AF. Floral dip:a simplified method for Agrobacterium-mediated transformation of Arabidopsis thaliana. Plant J,1998,16:735-743
23. Cohen P. Novel protein serine/threonine phosphatases:variety is the spice of life. Trends Biochem Sci, 1997,22:245
24. da Silva PM, Eastmond PJ, Hill LM, Smith AM, Rawsthorne S. Starch metabolism in developing embryos of oilseed rape. Planta, 1997,203:480-487
25. Dafny-Yelin M, Tzfira T, Vainstein A, Adam Z. Non-redundant functions of sHSP-CIs in acquired thermotolerance and their role in early seed development in Arabidopsis. Plant Mol Biol,2008,67:363-373
26. Davletova S, Schlauch K, Coutu J, Mittler R. The zinc-finger protein Zat12 plays a central role in reactive oxygen and abiotic stress signaling in Arabidopsis. Plant Physiol,2005,139:847-856
27. de Bouille P, Sotta B, Miginiac E, Merrien A. Hormones and pod development in oilseed rape (Brassica napus). Plant Physiol, 1989, 90:876-880
28. Dekkers BJ, Schuurmans JA, Smeekens SC. Glucose delays seed germination in Arabidopsis thaliana. Planta,2004,218:579-588
29. Deynze AV, Beversdorf W, Pauls K. Temperature effects on seed color in black-and yellow-seeded rapeseed. Can J Plant Sci, 1993,73:383-387
30. Endo M, Tsuchiya T, Hamada K, Kawamura S, Yano K, Ohshima M, Higashitani A, Watanabe M, Kawagishi-Kobayashi M. High temperatures cause male sterility in rice plants with transcriptional alterations during pollen development. Plant Cell Physiol,2009,50:1911-1922
31. Finkelstein R. Abscisic acid synthesis and response. Arabidopsis Book,2013, e0166
32. Fowler D, Downey R. Lipid and morphological changes in developing rapeseed, Brassica napus. Can JPlant Sci, 1970,50:233-247
33. Fucile G, Garcia C, Carlsson J, Sunnerhagen M, Christendat D. Structural and biochemical investigation of two Arabidopsis shikimate kinases:The heat-inducible isoform is thermostable. Protein Sci, 2011,20:1125-1136
34. Gan Y, Angadi S, Cutforth H, Potts D, Angadi V, McDonald C. Canola and mustard response to short periods of temperature and water stress at different developmental stages. Can J Plant Sci, 2004,84:697-704
35. Gao H, Brandizzi F, Benning C, Larkin RM. A membrane-tethered transcription factor defines a branch of the heat stress response in Arabidopsis thaliana. Proc Natl Acad Sci USA, 2008,105:16398-16403
36. Giorno F, Guerriero G, Baric S, Mariani C. Heat shock transcriptional factors in Malus domestica:identification, classification and expression analysis. BMC Genomics,2012,13:639
37. Giorno F, Wolters-Arts M, Mariani C, Rieu I. Ensuring reproduction at high temperatures: the heat stress response during anther and pollen development. Plants, 2013,2:489-506
38. Giraud E, Van Aken O, Ho LH, Whelan J. The transcription factor ABI4 is a regulator of mitochondrial retrograde expression of ALTERNATIVE OXIDASE1a. Plant Physiol,2009,150:1286-1296
39. Gray WM, Ostin A, Sandberg G, Romano CP, Estelle M. High temperature promotes auxin-mediated hypocotyl elongation in Arabidopsis. Proc Natl Acad Sci USA,1998, 95:7197-7202
40. Guan M, Li X, Guan C. Microarray analysis of differentially expressed genes between Brassica napus strains with high- and low-oleic acid contents. Plant Cell Rep,2012,31:929-943
41. Guan Q, Lu X, Zeng H, Zhang Y, Zhu J. Heat stress induction of miR398 triggers a regulatory loop that is critical for thermotolerance in Arabidopsis. Plant J, 2013,74: 840-851
42. Gulledge AA, Roberts AD, Vora H, Patel K, Loraine AE. Mining Arabidopsis thaliana RNA-seq data with Integrated Genome Browser reveals stress-induced alternative splicing of the putative splicing regulator SR45a. Am JBot, 2012, 99: 219-231
43. Guo FQ, Okamoto M, Crawford NM. Identification of a plant nitric oxide synthase gene involved in hormonal signaling. Science,2003,302:100-103
44. Guo J, Wu J, Ji Q, Wang C, Luo L, Yuan Y, Wang Y, Wang J. Genome-wide analysis of heat shock transcription factor families in rice and Arabidopsis. J Genet Genomics, 2008a,35:105-118
45. Guo L, Chen S, Liu K, Liu Y, Ni L, Zhang K, Zhang L. Isolation of heat shock factor HsfAla-binding sites in vivo revealed variations of heat shock elements in Arabidopsis thaliana. Plant Cell Physiol,2008b,49:1306-1315
46. Hall AE. Breeding for heat tolerance. Plant Breed Rev,1992,10:129-168
47. Herrero MP, Johnson R. High temperature stress and pollen viability of maize. Crop Sci,1980,20:796-800
48. Hong SW, Vierling E. Hsp101 is necessary for heat tolerance but dispensable for development and germination in the absence of stress. Plant J,2001,27:25-35
49. Hong SW, Vierling E. Mutants of Arabidopsis thaliana defective in the acquisition of tolerance to high temperature stress. Proc Natl Acad Sci USA,2000,97: 4392-4397
50. Hongo S, Sato K, Yokoyama R, Nishitani K. Demethylesterification of the primary wall by PECTIN METHYLESTERASE35 provides mechanical support to the Arabidopsis stem. Plant Cell,2012,24:2624-2634
51. Horton P, Nakai K. Better prediction of protein cellular localization sites with the k nearest neighbors classifier. Proc Int Conf Intell Syst Mol Biol.1997:147-152.
52. Hu C, Lin SY, Chi WT, Charng YY. Recent gene duplication and subfunctionalization produced a mitochondrial GrpE, the nucleotide exchange factor of the Hsp70 complex, specialized in thermotolerance to chronic heat stress in Arabidopsis. Plant Physiol,2012,158:747-758
53. Hua W, Li RJ, Zhan GM, Liu J, Li J, Wang XF, Liu GH, Wang HZ. Maternal control of seed oil content in Brassica napus: the role of silique wall photosynthesis. Plant J, 2012,69:432-444
54. Huang Y, Chen L, Wang L, Vijayan K, Phan S, Liu Z, Wan L, Ross A, Xiang D, Datla R. Probing the endosperm gene expression landscape in Brassica napus. BMC Genomics,2009,10:256
55. Ikeda M, Mitsuda N, Ohme-Takagi M. Arabidopsis HsfB1 and HsfB2b act as repressors of the expression of heat-inducible Hsfs but positively regulate the acquired thermo tolerance. Plant Physiol,2011,157:1243-1254
56. Jenkins ME, Suzuki TC, Mount DW. Evidence That heat and ultraviolet radiation activate a common stress-response program in plants that is alterd in the uvh6 mutant of Arabidopsis thaliana. Plant Physiol,1997,115:1351-1358
57. Johnson-Flanagan AM, Spencer MS. Ethylene production during development of mustard(Brassica juncea) and canola(Brassica napus) seed. Plant Physiol,1994, 106:601-606
58. Jonsson R. Breeding for improved oil and meal quality in rape(Brassica napus L.) and turnip rape(Brassica campestris L.). Hereditas,1978,87:205-218
59. Jungkunz I, Link K, Vogel F, Voll LM, Sonnewald S, Sonnewald U. AtHsp70-15-deficient Arabidopsis plants are characterized by reduced growth, a constitutive cytosolic protein response and enhanced resistance to TuMV. Plant J, 2011,66:983-995
60. Kanchiswamy CN, Muroi A, Maffei ME, Yoshioka H, Sawasaki T, Arimura G-i. Ca2+ -dependent protein kinases and their substrate HsfB2a are differently involved in the heat response signaling pathway in Arabidopsis. Plant Biotechnol,2010,27: 469-473
61. Kim M, Lee U, Small I, des Francs-Small CC, Vierling E. Mutations in an Arabidopsis mitochondrial transcription termination factor-related protein enhance thermotolerance in the absence of the major molecular chaperone HSP101. Plant Cell,2012,24:3349-3365
62. Kittock D, Turcotte E, Hofmann W. Estimation of heat tolerance improvement in recent American pima cotton cultivars. JAgron Crop Sci,1988,161:305-309
63. Ko CB, Woo YM, Lee DJ, Lee MC, Kim CS. Enhanced tolerance to heat stress in transgenic plants expressing the GASA4 gene. Plant Physiol Biochem,2007,45: 722-728
64. Krishna P, Gloor G. The Hsp90 family of proteins in Arabidopsis thaliana. Cell Stress Chaperones,2001,6:238-246
65. Kumar M, Busch W, Birke H, Kemmerling B, Nurnberger T, Schoffl F. Heat shock factors HsfB1 and HsfB2b are involved in the regulation of Pdf1.2 expression and pathogen resistance in Arabidopsis. Mol Plant,2009,2:152-165
66. Kuno N, M?ller SG, Shinomura T, Xu X, Chua NH, Furuya M. The novel MYB protein EARLY-PHYTOCHROME-RESPONSIVE1 is a component of a slave circadian oscillator in Arabidopsis. Plant Cell,2003,15:2476-2488
67. Kutcher H, Warland J, Brandt S. Temperature and precipitation effects on canola yields in Saskatchewan, Canada. Agric For Meteorol,2010,150:161-165
68. Kutuzov MA, Bennett N, Andreeva AV. Interaction of plant protein Ser/Thr phosphatase PP7 with calmodulin. Biochem Biophys Res Commun,2001,289: 634-640
69. Lamesch P, Berardini TZ, Li D, Swarbreck D, Wilks C, Sasidharan R, Muller R, Dreher K, Alexander DL, Garcia-Hernandez M. The Arabidopsis Information Resource (TAIR):improved gene annotation and new tools. Nucleic Acids Res,2012, 40:D1202-D1210
70. Larkindale J, Hall JD, Knight MR, Vierling E. Heat stress phenotypes of Arabidopsis mutants implicate multiple signaling pathways in the acquisition of thermotolerance. Plant Physiol,2005,138:882-897
71. Larkindale J, Knight MR. Protection against heat stress-induced oxidative damage in Arabidopsis involves calcium, abscisic acid, ethylene, and salicylic acid. Plant Physiol,2002,128:682-695
72. Larkindale J, Vierling E. Core genome responses involved in acclimation to high temperature. Plant Physiol,2008,146:748-761
73. Lee JH, Hiibel A, Schoffl F. Derepression of the activity of genetically engineered heat shock factor causes constitutive synthesis of heat shock proteins and increased thermotolerance in transgenic Arabidopsis. Plant J,1995,8:603-612
74. Lee JH, Schoffl F. An Hsp70 antisense gene affects the expression of HSP70/HSC70, the regulation of HSF, and the acquisition of thermotolerance in transgenic Arabidopsis thaliana. Mol Gen Genet, 1996,252:11-19
75. Lee S, Seo PJ, Lee HJ, Park CM. ANAC transcription factor NTL4 promotes reactive oxygen species production during drought-induced leaf senescence in Arabidopsis. Plant J,2012,70:831-844
76. Lee U, Rioflorido I, Hong SW, Larkindale J, Waters ER, Vierling E. The Arabidopsis ClpB/Hsp100 family of proteins: chaperones for stress and chloroplast development. Plant J,2007,49:115-127
77. Lee U, Wie C, Fernandez BO, Feelisch M, Vierling E. Modulation of nitrosative stress by S-nitrosoglutathione reductase is critical for thermotolerance and plant growth in Arabidopsis. Plant Cell,2008,20:786-802
78. Li GL, Chang H, Li B, Zhou W, Sun DY, Zhou RG. The roles of the atDjA2 and atDJA3 molecular chaperone proteins in improving thermotolerance of Arabidopsis thaliana seedlings. Plant Sci,2007,173:408-416
79. Li M, Berendzen KW, Schoeffl F. Promoter specificity and interactions between early and late Arabidopsis heat shock factors. Plant Mol Biol,2010,73:559-567
80. Li Z, Zhang L, Wang A, Xu X, Li J. Ectopic overexpression of SIHsfA3, a heat stress transcription factor from tomato, confers increased thermotolerance and salt hypersensitivity in germination in transgenic Arabidopsis. PLoS ONE,2013,8: e54880
81. Lillo C, Lea US, Ruoff P. Nutrient depletion as a key factor for manipulating gene expression and product formation in different branches of the flavonoid pathway. Plant Cell Environ,2008,31:587-601
82. Lin BL, Wang JS, Liu HC, Chen RW, Meyer Y, Barakat A, Delseny M. Genomic analysis of the Hsp70 superfamily in Arabidopsis thaliana. Cell Stress Chaperones, 2001,6:201-208
83. Lin YX, Jiang HY, Chu ZX, Tang XL, Zhu SW, Cheng BJ. Genome-wide identification, classification and analysis of heat shock transcription factor family in maize. BMC Genomics,2011,12:76
84. Lindquist S. The heat-shock response. Annu Rev Biochem,1986,55:1151-1191
85. Liu H, Charng Y. Acquired thermotolerance independent of heat shock factor Al (HsfAl), the master regulator of the heat stress response. Plant Signal Behav,2012, 7:547-550
86. Liu HC, Charng YY. Common and distinct functions of Arabidopsis class A1 and A2 heat shock factors in diverse abiotic stress responses and development. Plant Physiol, 2013,163:276-290
87. Liu HC, Liao HT, Charng YY. The role of class A1 heat shock factors (HSFA1s) in response to heat and other stresses in Arabidopsis. Plant Cell Environ,2011,34: 738-751
88. Liu HT, Gao F, Li GL, Han JL, De-Long L, Sun DY, Zhou RG. The calmodulin-binding protein kinase 3 is part of heat-shock signal transduction in Arabidopsis thaliana. Plant J,2008,55:760-773
89. Liu HT, Li GL, Chang H, Sun DY, Zhou RG, Li B. Calmodulin-binding protein phosphatase PP7 is involved in thermotolerance in Arabidopsis. Plant Cell Environ, 2007,30:156-164
90. Liu HT, Sun DY, Zhou RG. Ca2+ and AtCaM3 are involved in the expression of heat shock protein gene in Arabidopsis. Plant Cell Environ,2005,28:1276-1284
91. Liu J, Sun N, Liu M, Liu J, Du B, Wang X, Qi X. An autoregulatory loop controlling Arabidopsis HsfA2 expression: role of heat shock-induced alternative splicing. Plant Physiol,2013,162:512-521
92. Liu JG, Qin QL, Zhang Z, Peng RH, Xiong AS, Chen JM, Yao QH. OsHSF7 gene in rice, Oryza sativa L., encodes a transcription factor that functions as a high temperature receptive and responsive factor. BMB Rep,2009,42:16-21
93. Lohmann C, Eggers-Schumacher G, Wunderlich M, Schoffl F. Two different heat shock transcription factors regulate immediate early expression of stress genes in Arabidopsis. Mol Genet Genomics,2004,271:11-21
94. Lomascolo A, Uzan-Boukhris E, Sigoillot JC, Fine F. Rapeseed and sunflower meal: a review on biotechnology status and challenges. Appl Microbiol Biotechnol, 2012, 95:1105-1114
95. Martin T, Schmidt R, Altmann T, Frommer WB. Non-destructive assay systems for detection of β-glucuronidase activity in higher plants. Plant Mol Biol Reporter,1992, 10:37-46
96. Maruyama-Nakashita A, Nakamura Y, Tohge T, Saito K, Takahashi H. Arabidopsis SLIM1 is a central transcriptional regulator of plant sulfur response and metabolism. Plant Cell,2006,18:3235-3251
97. Meiri D, Tazat K, Cohen-Peer R, Farchi-Pisanty O, Aviezer-Hagai K, Avni A, Breiman A. Involvement of Arabidopsis ROF2 (FKBP65) in thermotolerance. Plant Mol Biol,2010,72:191-203
98. Miernyk JA. The J-domain proteins of Arabidopsis thaliana:an unexpectedly large and diverse family of chaperones. Cell Stress Chaperones,2001,6:209-218
99. Mishra SK, Tripp J, Winkelhaus S, Tschiersch B, Theres K, Nover L, Scharf KD. In the complex family of heat stress transcription factors, HSfA1 has a unique role as master regulator of thermotolerance in tomato. Gene Dev,2002,16:1555-1567
100. Mittler R, Finka A, Goloubinoff P. How do plants feel the heat? Trends Biochem Sci, 2012,37:118-125
101. Morrison M. Heat stress during reproduction in summer rape. Can J Bot,1993,71: 303-308
102. Morrison MJ, Stewart DW. Heat stress during flowering in summer Brassica. Crop Sci,2002,42:797-803
103. Mun JH, Kwon SJ, Yang TJ, Seol YJ, Jin M, Kim JA, Lim MH, Kim JS, Baek S, Choi BS. Genome-wide comparative analysis of the Brassica rapa gene space reveals genome shrinkage and differential loss of duplicated genes after whole genome triplication. Genome Biol,2009,10:R1ll
104. Murphy DJ, Cummins I. Biosynthesis of seed storage products during embryogenesis in rapeseed, Brassica napus. J Plant Physiol,1989,135:63-69
105.Nemhauser JL, Feldman LJ, Zambryski PC. Auxin and ETTIN in Arabidopsis gynoecium morphogenesis. Development,2000,127:3877-3888
106. Nesi N, Lucas MO, Auger B, Baron C, Lecureuil A, Guerche P, Kronenberger J, Lepiniec L, Debeaujon I, Renard M. The promoter of the Arabidopsis thaliana BAN gene is active in proanthocyanidin-accumulating cells of the Brassica napus seed coat. Plant Cell Rep,2009,28:601-617
107. Nishizawa A, Yabuta Y, Yoshida E, Maruta T, Yoshimura K, Shigeoka S. Arabidopsis heat shock transcription factor A2 as a key regulator in response to several types of environmental stress. Plant J,2006,48:535-547
108. Nishizawa-Yokoi A, Nosaka R, Hayashi H, Tainaka H, Maruta T, Tamoi M, Ikeda M, Ohme-Takagi M, Yoshimura K, Yabuta Y, Shigeoka S. HsfAld and HsfAle involved in the transcriptional regulation of HsfA2 function as key regulators for the Hsf signaling network in response to environmental stress. Plant Cell Physiol,2011,52: 933-945
109. Nishizawa-Yokoi A, Yoshida E, Yabuta Y, Shigeoka S. Analysis of the regulation of target genes by an Arabidopsis heat shock transcription factor, HsfA2. Biosci Biotechnol Biochem,2009,73:890-895
110. Noh YS, Amasino RM. Regulation of developmental senescence is conserved between Arabidopsis and Brassica napus. Plant Mol Biol,1999,41:195-206
111. Nour-Eldin HH, Andersen TG, Burow M, Madsen SR, J?rgensen ME, Olsen CE, Dreyer I, Hedrich R, Geiger D, Halkier BA. NRT/PTR transporters are essential for translocation of glucosinolate defence compounds to seeds. Nature,2012,488: 531-534
112.Nover L, Bharti K, Doering P, Mishra SK, Ganguli A, Scharf KD. Arabidopsis and the heat stress transcription factor world: How many heat stress transcription factors do we need? Cell Stress Chaperones,2001,6:177-189
113. Panikulangara TJ, Eggers-Schumacher G, Wunderlich M, Stransky H, Schoffl F. Galactinol synthasel. A novel heat shock factor target gene responsible for heat-induced synthesis of raffinose family oligosaccharides in Arabidopsis. Plant Physiol,2004,136:3148-3158
114. Parcy F, Valon C, Raynal M, Gaubier-Comella P, Delseny M, Giraudat J. Regulation of gene expression programs during Arabidopsis seed development:roles of the ABI3 locus and of endogenous abscisic acid. Plant Cell,1994,6:1567-1582
115. Park CH, Kim TW, Son SH, Hwang JY, Lee SC, Chang SC, Kim SH, Kim SW, Kim SK. Brassinosteroids control AtEXPA5 gene expression in Arabidopsis thaliana. Phytochemistry,2010,71:380-387
116. Park HY, Seok HY, Woo DH, Lee SY, Tarte VN, Lee EH, Lee CH, Moon YH. AtERF71/HRE2 transcription factor mediates osmotic stress response as well as hypoxia response in Arabidopsis. Biochem Biophys Res Commun,2011,414: 135-141
117. Park JH, Lee SY, Kim WY, Jung YJ, Chae HB, Jung HS, Kang CH, Shin MR, Kim SY, Su'udi M, Yun DJ, Lee KO, Kim MG, Lee SY. Heat-induced chaperone activity of serine/threonine protein phosphatase 5 enhances thermotolerance in Arabidopsis thaliana. New Phytol,2011,191:692-705
118. Paulsen GM. High temperature responses of crop plants. In: Boote KJ, Physiology and determination of crop yield. University of Minnesota: American Society of Agronomy,1994.365-389
119. Peet M, Sato S, Gardner R. Comparing heat stress effects on male-fertile and male-sterile tomatoes. Plant Cell Environ,1998,21:225-231
120. Peltonen-Sainio P, Jauhiainen L, Trnka M, Olesen JE, Calanca P, Eckersten H, Eitzinger J, Gobin A, Kersebaum KC, Kozyra J. Coincidence of variation in yield and climate in Europe. Agric Ecosyst Environ,2010, 139:483-489
121. Perez DE, Hoyer JS, Johnson AI, Moody ZR, Lopez J, Kaplinsky NJ. BOBBER1 is a noncanonical Arabidopsis small heat shock protein required for both development and thermotolerance. Plant Physiol,2009,151:241-252
122. Poschet G, Hannich B, Raab S, Jungkunz I, Klemens PA, Krueger S, Wic S, Neuhaus HE, Biittner M. A novel Arabidopsis vacuolar glucose exporter is involved in cellular sugar homeostasis and affects the composition of seed storage compounds. Plant Physiol,2011,157:1664-1676
123. Prandl R, Hinderhofer K, Eggers-Schumacher G, Schoffl F. HSF3, a new heat shock factor from Arabidopsis thaliana, derepresses the heat shock response and confers thermotolerance when overexpressed in transgenic plants. Mol Gen Genet, 1998,258: 269-278
124. Prasad BD, Goel S, Krishna P. In silico identification of carboxylate clamp type tetratricopeptide repeat proteins in Arabidopsis and rice as putative co-chaperones of Hsp90/Hsp70. PLoS ONE, 2010,5:e12761
125. Qu C, Fu F, Lu K, Zhang K, Wang R, Xu X, Wang M, Lu J, Wan H, Zhanglin T. Differential accumulation of phenolic compounds and expression of related genes in black- and yellow-seeded Brassica napus. J Exp Bot,2013,64:2885-2898
126. Queitsch C, Hong SW, Vierling E, Lindquist S. Heat shock protein 101 plays a crucial role in thermotolerance in Arabidopsis. Plant Cell,2000,12:479-492
127. Rasbery JM, Shan H, LeClair RJ, Norman M, Matsuda SP, Bartel B. Arabidopsis thaliana squalene epoxidase 1 is essential for root and seed development. JBiol Chem,2007,282:17002-17013
128.Rivero RM, Ruiz JM, Garcia PC, Lopez-Lefebre LR, Sanchez E, Romero L. Resistance to cold and heat stress:accumulation of phenolic compounds in tomato and watermelon plants. Plant Sci, 2001,160:315-321
129. Rizhsky L, Liang H, Mittler R. The combined effect of drought stress and heat shock on gene expression in tobacco. Plant Physiol,2002,130:1143-1151
130. Rizhsky L, Liang H, Shuman J, Shulaev V, Davletova S, Mittler R. When defense pathways collide. The response of Arabidopsis to a combination of drought and heat stress. Plant Physiol,2004,134:1683-1696
131. Romero HM, Berlett BS, Jensen PJ, Pell EJ, Tien M. Investigations into the role of the plastidial peptide methionine sulfoxide reductase in response to oxidative stress in Arabidopsis. Plant Physiol,2004,136:3784-3794
132. Rose J, Catala C, Urbanowicz B. Carbohydrate binding plant hydrolases which alter plant cell walls. USA Patent,0333223 Al.2010-12-30
133. Rowland O, Zheng H, Hepworth SR, Lam P, Jetter R, Kunst L. CER4 encodes an alcohol-forming fatty acyl-coenzyme A reductase involved in cuticular wax production in Arabidopsis. Plant Physiol,2006,142:866-877
134. Saini H, Sedgley M, Aspinall D. Effect of heat stress during floral development on pollen tube growth and ovary anatomy in wheat (Triticum aestivum L.). Funct Plant Biol,1983,10:137-144
135. Sakuma Y, Maruyama K, Osakabe Y, Qin F, Seki M, Shinozaki K, Yamaguchi-Shinozaki K. Functional analysis of an Arabidopsis transcription factor, DREB2A, involved in drought-responsive gene expression. Plant Cell,2006a, 18: 1292-1309
136. Sakuma Y, Maruyama K, Qin F, Osakabe Y, Shinozaki K, Yamaguchi-Shinozaki K. Dual function of an Arabidopsis transcription factor DREB2A in water-stress-responsive and heat-stress-responsive gene expression. Proc Natl Acad Sci USA,2006b,103:18822-18827
137. Samsel KA. The role of chloroplast-localized HSP21 in the stress responses of Arabidopsis thaliana. [Bachelors degree]. The University of Arizona: 2010
138. Sato S, Peet MM, Thomas JF. Determining critical pre- and post-anthesis periods and physiological processes in Lycopersicon esculentum Mill, exposed to moderately elevated temperatures. JExp Bot,2002,53:1187-1195
139. Scharf KD, Berberich T, Ebersberger I, Nover L. The plant heat stress transcription factor (Hsf) family: structure, function and evolution. Biochim Biophys Acta,2011, 1819:104-119
140. Scharf KD, Heider H, Hohfeld I, Lyck R, Schmidt E, Nover L. The tomato Hsf system: HsfA2 needs interaction with HsfAl for efficient nuclear import and may be localized in cytoplasmic heat stress granules. Mol Cell Biol, 1998,18:2240-2251
141. Scharf KD, Siddique M, Vierling E. The expanding family of Arabidopsis thaliana small heat stress proteins and a new family of proteins containing α-crystallin domains (Acd proteins). Cell Stress Chaperones,2001,6:225-237
142.Schlesinger MJ. Heat shock proteins. JBiol Chem,1990,265:12111-12114
143. Schramm F, Ganguli A, Kiehlmann E, Englich G, Walch D, von Koskull-Doring P. The heat stress transcription factor HsfA2 serves as a regulatory amplifier of a subset of genes in the heat stress response in Arabidopsis. Plant Mol Biol,2006,60: 759-772
144. Schramm F, Larkindale J, Kiehlmann E, Ganguli A, Englich G, Vierling E, von Koskull-Doering P. A cascade of transcription factor DREB2A and heat stress transcription factor HsfA3 regulates the heat stress response of Arabidopsis. Plant J, 2008,53:264-274
145. Shonnard GC, Gepts P. Genetics of heat tolerance during reproductive development in common bean. Crop Sci,1994,34:1168-1175
146. Siddique M, Gernhard S, von Koskull-Doering P, Vierling E, Scharf KD. The plant sHSP superfamily: five new members in Arabidopsis thaliana with unexpected properties. Cell Stress Chaperones,2008,13:183-197
147. S?nderby IE, Geu-Flores F, Halkier BA. Biosynthesis of glucosinolates-gene discovery and beyond. Trends Plant Sci,2010,15:283-290
148. Su PH, Li HM. Arabidopsis stromal 70-kD heat shock proteins are essential for plant development and important for thermotolerance of germinating seeds. Plant Physiol, 2008,146:1231-1241
149. Subramanian A, Tamayo P, Mootha VK, Mukherjee S, Ebert BL, Gillette MA, Paulovich A, Pomeroy SL, Golub TR, Lander ES. Gene set enrichment analysis:a knowledge-based approach for interpreting genome-wide expression profiles. Proc Natl Acad Sci USA,2005,102:15545-15550
150. Sugio A, Dreos R, Aparicio F, Maule AJ. The cytosolic protein response as a subcomponent of the wider heat shock response in Arabidopsis. Plant Cell,2009,21: 642-654
151. Suzuki N, Bajad S, Shuman J, Shulaev V, Mittler R. The transcriptional co-activator MBFlc is a key regulator of thermo tolerance in Arabidopsis thaliana. J Biol Chem, 2008,283:9269-9275
152. Suzuki N, Rizhsky L, Liang HJ, Shuman J, Shulaev V, Mittler R. Enhanced tolerance to environmental stress in transgenic plants expressing the transcriptional coactivator multiprotein bridging factor 1c. Plant Physiol,2005,139:1313-1322
153. Suzuki N, Sejima H, Tam R, Schlauch K, Mittler R. Identification of the MBF1 heat-response regulon of Arabidopsis thaliana. Plant J,2011,66:844-851
154. Swindell WR, Huebner M, Weber AP. Transcriptional profiling of Arabidopsis heat shock proteins and transcription factors reveals extensive overlap between heat and non-heat stress response pathways. BMC Genomics,2007,8:125
155. Tanabe N, Yoshimura K, Kimura A, Yabuta Y, Shigeoka S. Differential expression of alternatively spliced mRNAs of Arabidopsis SR protein homologs, atSR30 and atSR45a, in response to environmental stress. Plant Cell Physiol,2007,48: 1036-1049
156. Tayo T, Morgan D. Quantitative analysis of the growth, development and distribution of flowers and pods in oilseed rape (Brassica napus L.). JAgric Sci, Cambridge,1975,85:103-110
157. Thomas J, Palusa SG, Prasad KV, Ali GS, Surabhi GK, Ben-Hur A, Abdel-Ghany SE, Reddy AS. Identification of an intronic splicing regulatory element involved in auto-regulation of alternative splicing of SCL33 pre-mRNA. Plant J,2012,72: 935-946
158. Trenberth KE, Jones PD., Ambenje P, Bojariu R, Easterling D, Tank AK, Parker D, Rahimzadeh F, Renwick JA, Rusticucci M, Soden B and Zhai P. Observations: surface and atmospheric climate change. In: Solomon, S., Qin, D., Manning, M., Chen, Z., Marquis, M., Averyt, K.B., Tignor, M.,Miller, H.L. eds., Climate change 2007:the physical science basis. Contribution of working group I to the fourth assessment report of the Intergovernmental Panel on Climate Change. Cambridge, United Kingdom and New York, NY, USA: Cambridge University Press,2007.
159. Trick M, Cheung F, Drou N, Fraser F, Lobenhofer EK, Hurban P, Magusin A, Town CD, Bancroft I. A newly-developed community microarray resource for transcriptome profiling in Brassica species enables the confirmation of Brassica-specific expressed sequences. BMC Plant Biol,2009,9:50
160. Tsutsui T, Kato W, Asada Y, Sako K, Sato T, Sonoda Y, Kidokoro S, Yamaguchi-Shinozaki K, Tamaoki M, Arakawa K. DEAR1, a transcriptional repressor of DREB protein that mediates plant defense and freezing stress responses in Arabidopsis. J Plant Res,2009,122:633-643
161. Vivian-Smith A, Koltunow AM. Genetic analysis of growth-regulator-induced parthenocarpy in Arabidopsis. Plant Physiol,1999,121:437-452
162. Vogel JT, Zarka DG, Van Buskirk HA, Fowler SG, Thomashow MF. Roles of the CBF2 and ZAT12 transcription factors in configuring the low temperature transcriptome of Arabidopsis. Plant J,2005,41:195-211
163. Volkov RA, Panchuk Ⅱ, Mullineaux PM, Schoeffl F. Heat stress-induced H2O2 is required for effective expression of heat shock genes in Arabidopsis. Plant Mol Biol, 2006,61:733-746
164. Wang X, Xu L, Zheng L. Cloning and expression of phosphatidylcholine-hydrolyzing phospholipase D from Ricinus communis L. J Biol Chem,1994,269:20312-20317
165. Watanabe N, Lam E. Arabidopsis Bax inhibitor-1 functions as an attenuator of biotic and abiotic types of cell death. Plant J,2006,45:884-894
166. Whitecross M, Armstrong D. Environmental effects on epicuticular waxes of Brassica napus L. Aust J Bot,1972,20:87-95
167. Wingenter K, Schulz A, Wormit A, Wic S, Trentmann O, Hoermiller II, Heyer AG, Marten I, Hedrich R, Neuhaus HE. Increased activity of the vacuolar monosaccharide transporter TMT1 alters cellular sugar partitioning, sugar signaling, and seed yield in Arabidopsis. Plant Physiol,2010,154:665-677
168. Wu C. Heat shock transcription factors:structure and regulation. Annu Rev Cell Dev Biol,1995,11:441-469
169. Wu CC, Singh P, Chen MC, Zimmerli L. L-Glutamine inhibits beta-aminobutyric acid-induced stress resistance and priming in Arabidopsis. JExp Bot,2010,61: 995-1002
170. Wu SJ, Wang LC, Yeh CH, Lu CA, Wu SJ. Isolation and characterization of the Arabidopsis heat-intolerant 2 (hit2) mutant reveal the essential role of the nuclear export receptor EXPORTTN1A (XPO1A) in plant heat tolerance. New Phytol,2010, 186:833-842
171. Wu TY, Juan YT, Hsu YH, Wu SH, Liao HT, Fung RW, Charng YY. Interplay between heat shock proteins, HSP101 and HSA32, prolongs heat acclimation memory posttranscriptionally in Arabidopsis. Plant Physiol,2013,161:2075-2084
172. Wunderlich M, Werr W, Schoffl F. Generation of dominant-negative effects on the heat shock response in Arabidopsis thaliana by transgenic expression of a chimaeric HSF1 protein fusion construct. Plant J,2003,35:442-451
173. Xin H, Zhang H, Chen L, Li X, Lian Q, Yuan X, Hu X, Cao L, He X, Yi M. Cloning and characterization of HsfA2 from Lily (Lilium longiflorum). Plant Cell Rep,2010, 29:875-885
174. Xing Y, Jia W, Zhang J. AtMKK1 and AtMPK6 are involved in abscisic acid and sugar signaling in Arabidopsis seed germination. Plant Mol Biol,2009,70:725-736
175. Xu X, Song H, Zhou Z, Shi N, Ying Q, Wang H. Functional characterization of AtHsp90.3 in Saccharomyces cerevisiae and Arabidopsis thaliana under heat stress. Biotechnol Lett,2010,32:979-987
176. Xuan Y, Zhou S, Wang L, Cheng Y, Zhao L. Nitric oxide functions as a signal and acts upstream of AtCaM3 in thermotolerance in Arabidopsis seedlings. Plant Physiol, 2010,153:1895-1906
177. Yamada K, Fukao Y, Hayashi M, Fukazawa M, Suzuki I, Nishimura M. Cytosolic HSP90 regulates the heat shock response that is responsible for heat acclimation in Arabidopsis thaliana. J Biol Chem,2007,282:37794-37804
178. Yamakawa H, Hakata M. Atlas of rice grain filling-related metabolism under high temperature:joint analysis of metabolome and transcriptome demonstrated inhibition of starch accumulation and induction of amino acid accumulation. Plant Cell Physiol, 2010,51:795-809
179. Yamakawa H, Hirose T, Kuroda M, Yamaguchi T. Comprehensive expression profiling of rice grain filling-related genes under high temperature using DNA microarray. Plant Physiol,2007,144:258-277
180. Yamamoto Y, Shimada Y, Kimura M, Manabe K, Sekine Y, Matsui M, Ryuto H, Fukunishi N, Abe T, Yoshida S. Global classification of transcriptional responses to light stress in Arabidopsis thaliana. Endocytobiosis Cell Res,2004,15:438-452
181. Yang C Y, Hsu FC, Li JP, Wang NN, Shih MC. The AP2/ERF transcription factor AtERF73/HRE1 modulates ethylene responses during hypoxia in Arabidopsis. Plant Physiol,2011,156:202-212
182. Yang K, Lim CJ, Hong JK, Park CY, Cheong YH, Chung WS, Lee KO, Lee S Y, Cho MJ, Lim CO. Identification of cell wall genes modified by a permissive high temperature in Chinese cabbage. Plant Sci,2006,171:175-182
183. Yang YW, Lai KN, Tai PY, Li WH. Rates of nucleotide substitution in angiosperm mitochondrial DNA sequences and dates of divergence between Brassica and other angiosperm lineages. J Mol Evol,1999,48:597-604
184. Yokotani N, Ichikawa T, Kondou Y, Matsui M, Hirochika H, Iwabuchi M, Oda K. Expression of rice heat stress transcription factor OsHsfA2e enhances tolerance to environmental stresses in transgenic Arabidopsis. Planta,2008,227:957-967
185. Yoo CY, Miura K, Jin JB, Lee J, Park HC, Salt DE, Yun DJ, Bressan RA, Hasegawa PM. SIZ1 small ubiquitin-like modifier E3 ligase facilitates basal thermotolerance in Arabidopsis independent of salicylic acid. Plant Physiol,2006,142:1548-1558
186. Yoo SD, Cho YH, Sheen J. Arabidopsis mesophyll protoplasts:a versatile cell system for transient gene expression analysis. Nat Protoc,2007,2:1565-1572
187. Yoshida T, Sakuma Y, Todaka D, Maruyama K, Qin F, Mizoi J, Kidokoro S, Fujita Y, Shinozaki K, Yamaguchi-Shinozaki K. Functional analysis of an Arabidopsis heat-shock transcription factor HsfA3 in the transcriptional cascade downstream of the DREB2A stress-regulatory system. Biochem Biophys Res Commun,2008,368: 515-521
188. Yoshimura K, Mori T, Yokoyama K, Koike Y, Tanabe N, Sato N, Takahashi H, Maruta T, Shigeoka S. Identification of alternative splicing events regulated by an Arabidopsis serine/arginine-like protein, atSR45a, in response to high-light stress using a tiling array. Plant Cell Physiol,2011,52:1786-1805
189. Young LW, Wilen RW, Bonham-Smith PC. High temperature stress of Brassica napus during flowering reduces micro- and megagametophyte fertility, induces fruit abortion, and disrupts seed production. JExp Bot,2004,55:485-495
190. Yu HD, Yang XF, Chen ST, Wang YT, Li JK, Shen Q, Liu XL, Guo FQ. Downregulation of chloroplast RPS1 negatively modulates nuclear heat-responsive expression of HsfA2 and its target genes in Arabidopsis. Plos Genet,2012a,8: e1002669
191. Yu X, Wang H, Lu Y, de Ruiter M, Cariaso M, Prins M, van Tunen A, He Y. Identification of conserved and novel microRNAs that are responsive to heat stress in Brassica rapa. J Exp Bot, 2012b,63:1025-1038
192. Zanewich KP, Rood SB. Distribution of endogenous gibberellins in vegetative and reproductive organs of Brassica. J Plant Growth Regul,1993,12:41-46
193. Zelisko A, Garcia-Lorenzo M, Jackowski G, Jansson S, Funk C. AtFtsH6 is involved in the degradation of the light-harvesting complex II during high-light acclimation and senescence. Proc Natl Acad Sci USA,2005,102:13699-13704
194. Zhang H, Yang J, Chen Y, Mao X, Wang Z, Li C. Identification and expression analysis of the heat shock transcription factor (HSF) gene family in Populus trichocarpa. Plant Omics.2013,6:415-424
195. Zhang LR, Li YS, Xing D, Gao CJ. Characterization of mitochondrial dynamics and subcellular localization of ROS reveal that HsfA2 alleviates oxidative damage caused by heat stress in Arabidopsis. J Exp Bot,2009a,60:2073-2091
196. Zhang W, Zhou RG, Gao YJ, Zheng SZ, Xu P, Zhang SQ, Sun DY. Molecular and genetic evidence for the key role of AtCaM3 in heat-shock signal transduction in Arabidopsis. Plant Physiol,2009b,149:1773-1784
197. Zhou S, Zhang P, Jing Z, Shi J. Genome-wide identification and analysis of heat shock transcription factor family in cucumber (Cucumis sativus L.). Plant Omics. 2013,6:449-455
198. Zhou W, Zhou T, Li MX, Zhao CL, Jia N, Wang XX, Sun YZ, Li GL, Xu M, Zhou RG. The Arabidopsis J-protein AtDjBl facilitates thermotolerance by protecting cells against heat-induced oxidative damage. New Phytol, 2012,194:364-378
199. Zhu G, Liu Y, Ye N, Liu R, Zhang J. Involvement of the abscisic acid catabolic gene CYP707A2 in the glucose-induced delay in seed germination and post-germination growth of Arabidopsis. Physiol Plant,2011,143:375-384
200. Zhu Y, Cao Z, Xu F, Huang Y, Chen M, Guo W, Zhou W, Zhu J, Meng J, Zou J. Analysis of gene expression profiles of two near-isogenic lines differing at a QTL region affecting oil content at high temperatures during seed maturation in oilseed rape(Brassica napus L.). Theor Appl Genet,2012,124:515-531
201. Zhu Y, Wang Z, Jing Y, Wang LL, Liu X, Liu YX, Deng X. Ectopic over-expression of BhHsfl, a heat shock factor from the resurrection plant Boea hygrometrica, leads to increased thermotolerance and retarded growth in transgenic Arabidopsis and tobacco. Plant Mol Biol,2009a,71:451-467
202. Zimmerli L, Hou BH, Tsai CH, Jakab G, Mauch-Mani B, Somerville S. The xenobiotic beta-aminobutyric acid enhances Arabidopsis thermotolerance. Plant J, 2008,53:144-156
2. 王汉中,华玮,刘静,刘贵华,王新发,杨庆.油菜热激蛋白基因HSP17.8及其应用.中国专利,101544983.2009-09-30
3. 张树杰,王汉中.我国油菜生产应对气候变化的对策和措施分析.中国油料作物学报,2012,34：114-122
4. Agarwal M, Katiyar-Agarwal S, Sahi C, Gallie DR, Grover A. Arabidopsis thaliana Hsp100 proteins:kith and kin. Cell Stress Chaperones,2001,6:219
5. Aksouh-Harradj N, Campbell L, Mailer R. Canola response to high and moderately high temperature stresses during seed maturation. Can J Plant Sci,2006,86: 967-980
6. Anderson LE. Hoyer's solution as a rapid permanent mounting medium for bryophytes. Bryologist,1954,57:242-244
7. Angadi S, Cutforth H, Miller P, McConkey B, Entz M, Brandt S, Volkmar K. Response of three Brassica species to high temperature stress during reproductive growth. Can J Plant Sci,2000,80:693-701
8. Baniwal SK, Chan KY, Scharf K-D, Nover L. Role of heat stress transcription factor HsfA5 as specific repressor of HsfA4. J Biol Chem,2007,282:3605-3613
9. Battisti DS, Naylor RL. Historical warnings of future food insecurity with unprecedented seasonal heat. Science,2009,323:240-244
10. Bharti K, von Ko skull-Dor ing P, Bharti S, Kumar P, Tintschl-Korbitzer A, Treuter E, Nover L. Tomato heat stress transcription factor HsfB1 represents a novel type of general transcription coactivator with a histone-like motif interacting with the plant CREB binding protein ortholog HAC1. Plant Cell,2004,16:1521-1535
11. Busch W, Wunderlich M, Schoffl F. Identification of novel heat shock factor-dependent genes and biochemical pathways in Arabidopsis thaliana. Plant J, 2004,41:1-14
12. Charng YY, Liu HC, Liu NY, Chi WT, Wang CN, Chang SH, Wang TT. A heat-inducible transcription factor, HsfA2, is required for extension of acquired thermotolerance in Arabidopsis. Plant Physiol,2007,143:251-262
13. Charng YY, Liu HC, Liu NY, Hsu FC, Ko SS. Arabidopsis Hsa32, a novel heat shock protein, is essential for acquired thermotolerance during long recovery after acclimation. Plant Physiol,2006,140:1297-1305
14. Chauhan H, Khurana N, Agarwal P, Khurana JP, Khurana P. A seed preferential heat shock transcription factor from wheat provides abiotic stress tolerance and yield enhancement in transgenic Arabidopsis under heat stress environment. PLoS ONE, 2013,8:e79577
15. Chauhan H, Khurana N, Tyagi AK, Khurana JP, Khurana P. Identification and characterization of high temperature stress responsive genes in bread wheat (Triticum aestivum L.) and their regulation at various stages of development. Plant Mol Biol,2011,75:35-51
16. Chen G, Bi YR, Li N. EGY1 encodes a membrane-associated and ATP-independent metalloprotease that is required for chloroplast development. Plant J,2005,41: 364-3751
17. Chen H, Hwang JE, Lim CJ, Kim DY, Lee SY, Lim CO. Arabidopsis DREB2C functions as a transcriptional activator of HsfA3 during the heat stress response. Biochem Biophys Res Commun,2010,401:238-244
18. Chen X, Ye C, Lii H, Xu M, Li W, Zhang L, Wang C, Luo S, Zhu B. Cloning of GmHSFAl gene and its overexpression leading to enhancement of heat tolerance in transgenic soybean. Yi Chuan,2006,28:1411-1420
19. Clarke SM, Cristescu SM, Miersch O, Harren FJM, Wasternack C, Mur LAJ. Jasmonates act with salicylic acid to confer basal thermotolerance in Arabidopsis thaliana. New Phytol, 2009,182:175-187
20. Clarke SM, Mur LAJ, Wood JE, Scott IM. Salicylic acid dependent signaling promotes basal thermotolerance but is not essential for acquired thermotolerance in Arabidopsis thaliana. Plant J,2004,38:432-447
21. Claros MG, Vincens P. Computational method to predict mitochondrially imported proteins and their targeting sequences. Eur J Biochem, 1996,241:779-786
22. Clough SJ, Bent AF. Floral dip:a simplified method for Agrobacterium-mediated transformation of Arabidopsis thaliana. Plant J,1998,16:735-743
23. Cohen P. Novel protein serine/threonine phosphatases:variety is the spice of life. Trends Biochem Sci, 1997,22:245
24. da Silva PM, Eastmond PJ, Hill LM, Smith AM, Rawsthorne S. Starch metabolism in developing embryos of oilseed rape. Planta, 1997,203:480-487
25. Dafny-Yelin M, Tzfira T, Vainstein A, Adam Z. Non-redundant functions of sHSP-CIs in acquired thermotolerance and their role in early seed development in Arabidopsis. Plant Mol Biol,2008,67:363-373
26. Davletova S, Schlauch K, Coutu J, Mittler R. The zinc-finger protein Zat12 plays a central role in reactive oxygen and abiotic stress signaling in Arabidopsis. Plant Physiol,2005,139:847-856
27. de Bouille P, Sotta B, Miginiac E, Merrien A. Hormones and pod development in oilseed rape (Brassica napus). Plant Physiol, 1989, 90:876-880
28. Dekkers BJ, Schuurmans JA, Smeekens SC. Glucose delays seed germination in Arabidopsis thaliana. Planta,2004,218:579-588
29. Deynze AV, Beversdorf W, Pauls K. Temperature effects on seed color in black-and yellow-seeded rapeseed. Can J Plant Sci, 1993,73:383-387
30. Endo M, Tsuchiya T, Hamada K, Kawamura S, Yano K, Ohshima M, Higashitani A, Watanabe M, Kawagishi-Kobayashi M. High temperatures cause male sterility in rice plants with transcriptional alterations during pollen development. Plant Cell Physiol,2009,50:1911-1922
31. Finkelstein R. Abscisic acid synthesis and response. Arabidopsis Book,2013, e0166
32. Fowler D, Downey R. Lipid and morphological changes in developing rapeseed, Brassica napus. Can JPlant Sci, 1970,50:233-247
33. Fucile G, Garcia C, Carlsson J, Sunnerhagen M, Christendat D. Structural and biochemical investigation of two Arabidopsis shikimate kinases:The heat-inducible isoform is thermostable. Protein Sci, 2011,20:1125-1136
34. Gan Y, Angadi S, Cutforth H, Potts D, Angadi V, McDonald C. Canola and mustard response to short periods of temperature and water stress at different developmental stages. Can J Plant Sci, 2004,84:697-704
35. Gao H, Brandizzi F, Benning C, Larkin RM. A membrane-tethered transcription factor defines a branch of the heat stress response in Arabidopsis thaliana. Proc Natl Acad Sci USA, 2008,105:16398-16403
36. Giorno F, Guerriero G, Baric S, Mariani C. Heat shock transcriptional factors in Malus domestica:identification, classification and expression analysis. BMC Genomics,2012,13:639
37. Giorno F, Wolters-Arts M, Mariani C, Rieu I. Ensuring reproduction at high temperatures: the heat stress response during anther and pollen development. Plants, 2013,2:489-506
38. Giraud E, Van Aken O, Ho LH, Whelan J. The transcription factor ABI4 is a regulator of mitochondrial retrograde expression of ALTERNATIVE OXIDASE1a. Plant Physiol,2009,150:1286-1296
39. Gray WM, Ostin A, Sandberg G, Romano CP, Estelle M. High temperature promotes auxin-mediated hypocotyl elongation in Arabidopsis. Proc Natl Acad Sci USA,1998, 95:7197-7202
40. Guan M, Li X, Guan C. Microarray analysis of differentially expressed genes between Brassica napus strains with high- and low-oleic acid contents. Plant Cell Rep,2012,31:929-943
41. Guan Q, Lu X, Zeng H, Zhang Y, Zhu J. Heat stress induction of miR398 triggers a regulatory loop that is critical for thermotolerance in Arabidopsis. Plant J, 2013,74: 840-851
42. Gulledge AA, Roberts AD, Vora H, Patel K, Loraine AE. Mining Arabidopsis thaliana RNA-seq data with Integrated Genome Browser reveals stress-induced alternative splicing of the putative splicing regulator SR45a. Am JBot, 2012, 99: 219-231
43. Guo FQ, Okamoto M, Crawford NM. Identification of a plant nitric oxide synthase gene involved in hormonal signaling. Science,2003,302:100-103
44. Guo J, Wu J, Ji Q, Wang C, Luo L, Yuan Y, Wang Y, Wang J. Genome-wide analysis of heat shock transcription factor families in rice and Arabidopsis. J Genet Genomics, 2008a,35:105-118
45. Guo L, Chen S, Liu K, Liu Y, Ni L, Zhang K, Zhang L. Isolation of heat shock factor HsfAla-binding sites in vivo revealed variations of heat shock elements in Arabidopsis thaliana. Plant Cell Physiol,2008b,49:1306-1315
46. Hall AE. Breeding for heat tolerance. Plant Breed Rev,1992,10:129-168
47. Herrero MP, Johnson R. High temperature stress and pollen viability of maize. Crop Sci,1980,20:796-800
48. Hong SW, Vierling E. Hsp101 is necessary for heat tolerance but dispensable for development and germination in the absence of stress. Plant J,2001,27:25-35
49. Hong SW, Vierling E. Mutants of Arabidopsis thaliana defective in the acquisition of tolerance to high temperature stress. Proc Natl Acad Sci USA,2000,97: 4392-4397
50. Hongo S, Sato K, Yokoyama R, Nishitani K. Demethylesterification of the primary wall by PECTIN METHYLESTERASE35 provides mechanical support to the Arabidopsis stem. Plant Cell,2012,24:2624-2634
51. Horton P, Nakai K. Better prediction of protein cellular localization sites with the k nearest neighbors classifier. Proc Int Conf Intell Syst Mol Biol.1997:147-152.
52. Hu C, Lin SY, Chi WT, Charng YY. Recent gene duplication and subfunctionalization produced a mitochondrial GrpE, the nucleotide exchange factor of the Hsp70 complex, specialized in thermotolerance to chronic heat stress in Arabidopsis. Plant Physiol,2012,158:747-758
53. Hua W, Li RJ, Zhan GM, Liu J, Li J, Wang XF, Liu GH, Wang HZ. Maternal control of seed oil content in Brassica napus: the role of silique wall photosynthesis. Plant J, 2012,69:432-444
54. Huang Y, Chen L, Wang L, Vijayan K, Phan S, Liu Z, Wan L, Ross A, Xiang D, Datla R. Probing the endosperm gene expression landscape in Brassica napus. BMC Genomics,2009,10:256
55. Ikeda M, Mitsuda N, Ohme-Takagi M. Arabidopsis HsfB1 and HsfB2b act as repressors of the expression of heat-inducible Hsfs but positively regulate the acquired thermo tolerance. Plant Physiol,2011,157:1243-1254
56. Jenkins ME, Suzuki TC, Mount DW. Evidence That heat and ultraviolet radiation activate a common stress-response program in plants that is alterd in the uvh6 mutant of Arabidopsis thaliana. Plant Physiol,1997,115:1351-1358
57. Johnson-Flanagan AM, Spencer MS. Ethylene production during development of mustard(Brassica juncea) and canola(Brassica napus) seed. Plant Physiol,1994, 106:601-606
58. Jonsson R. Breeding for improved oil and meal quality in rape(Brassica napus L.) and turnip rape(Brassica campestris L.). Hereditas,1978,87:205-218
59. Jungkunz I, Link K, Vogel F, Voll LM, Sonnewald S, Sonnewald U. AtHsp70-15-deficient Arabidopsis plants are characterized by reduced growth, a constitutive cytosolic protein response and enhanced resistance to TuMV. Plant J, 2011,66:983-995
60. Kanchiswamy CN, Muroi A, Maffei ME, Yoshioka H, Sawasaki T, Arimura G-i. Ca2+ -dependent protein kinases and their substrate HsfB2a are differently involved in the heat response signaling pathway in Arabidopsis. Plant Biotechnol,2010,27: 469-473
61. Kim M, Lee U, Small I, des Francs-Small CC, Vierling E. Mutations in an Arabidopsis mitochondrial transcription termination factor-related protein enhance thermotolerance in the absence of the major molecular chaperone HSP101. Plant Cell,2012,24:3349-3365
62. Kittock D, Turcotte E, Hofmann W. Estimation of heat tolerance improvement in recent American pima cotton cultivars. JAgron Crop Sci,1988,161:305-309
63. Ko CB, Woo YM, Lee DJ, Lee MC, Kim CS. Enhanced tolerance to heat stress in transgenic plants expressing the GASA4 gene. Plant Physiol Biochem,2007,45: 722-728
64. Krishna P, Gloor G. The Hsp90 family of proteins in Arabidopsis thaliana. Cell Stress Chaperones,2001,6:238-246
65. Kumar M, Busch W, Birke H, Kemmerling B, Nurnberger T, Schoffl F. Heat shock factors HsfB1 and HsfB2b are involved in the regulation of Pdf1.2 expression and pathogen resistance in Arabidopsis. Mol Plant,2009,2:152-165
66. Kuno N, M?ller SG, Shinomura T, Xu X, Chua NH, Furuya M. The novel MYB protein EARLY-PHYTOCHROME-RESPONSIVE1 is a component of a slave circadian oscillator in Arabidopsis. Plant Cell,2003,15:2476-2488
67. Kutcher H, Warland J, Brandt S. Temperature and precipitation effects on canola yields in Saskatchewan, Canada. Agric For Meteorol,2010,150:161-165
68. Kutuzov MA, Bennett N, Andreeva AV. Interaction of plant protein Ser/Thr phosphatase PP7 with calmodulin. Biochem Biophys Res Commun,2001,289: 634-640
69. Lamesch P, Berardini TZ, Li D, Swarbreck D, Wilks C, Sasidharan R, Muller R, Dreher K, Alexander DL, Garcia-Hernandez M. The Arabidopsis Information Resource (TAIR):improved gene annotation and new tools. Nucleic Acids Res,2012, 40:D1202-D1210
70. Larkindale J, Hall JD, Knight MR, Vierling E. Heat stress phenotypes of Arabidopsis mutants implicate multiple signaling pathways in the acquisition of thermotolerance. Plant Physiol,2005,138:882-897
71. Larkindale J, Knight MR. Protection against heat stress-induced oxidative damage in Arabidopsis involves calcium, abscisic acid, ethylene, and salicylic acid. Plant Physiol,2002,128:682-695
72. Larkindale J, Vierling E. Core genome responses involved in acclimation to high temperature. Plant Physiol,2008,146:748-761
73. Lee JH, Hiibel A, Schoffl F. Derepression of the activity of genetically engineered heat shock factor causes constitutive synthesis of heat shock proteins and increased thermotolerance in transgenic Arabidopsis. Plant J,1995,8:603-612
74. Lee JH, Schoffl F. An Hsp70 antisense gene affects the expression of HSP70/HSC70, the regulation of HSF, and the acquisition of thermotolerance in transgenic Arabidopsis thaliana. Mol Gen Genet, 1996,252:11-19
75. Lee S, Seo PJ, Lee HJ, Park CM. ANAC transcription factor NTL4 promotes reactive oxygen species production during drought-induced leaf senescence in Arabidopsis. Plant J,2012,70:831-844
76. Lee U, Rioflorido I, Hong SW, Larkindale J, Waters ER, Vierling E. The Arabidopsis ClpB/Hsp100 family of proteins: chaperones for stress and chloroplast development. Plant J,2007,49:115-127
77. Lee U, Wie C, Fernandez BO, Feelisch M, Vierling E. Modulation of nitrosative stress by S-nitrosoglutathione reductase is critical for thermotolerance and plant growth in Arabidopsis. Plant Cell,2008,20:786-802
78. Li GL, Chang H, Li B, Zhou W, Sun DY, Zhou RG. The roles of the atDjA2 and atDJA3 molecular chaperone proteins in improving thermotolerance of Arabidopsis thaliana seedlings. Plant Sci,2007,173:408-416
79. Li M, Berendzen KW, Schoeffl F. Promoter specificity and interactions between early and late Arabidopsis heat shock factors. Plant Mol Biol,2010,73:559-567
80. Li Z, Zhang L, Wang A, Xu X, Li J. Ectopic overexpression of SIHsfA3, a heat stress transcription factor from tomato, confers increased thermotolerance and salt hypersensitivity in germination in transgenic Arabidopsis. PLoS ONE,2013,8: e54880
81. Lillo C, Lea US, Ruoff P. Nutrient depletion as a key factor for manipulating gene expression and product formation in different branches of the flavonoid pathway. Plant Cell Environ,2008,31:587-601
82. Lin BL, Wang JS, Liu HC, Chen RW, Meyer Y, Barakat A, Delseny M. Genomic analysis of the Hsp70 superfamily in Arabidopsis thaliana. Cell Stress Chaperones, 2001,6:201-208
83. Lin YX, Jiang HY, Chu ZX, Tang XL, Zhu SW, Cheng BJ. Genome-wide identification, classification and analysis of heat shock transcription factor family in maize. BMC Genomics,2011,12:76
84. Lindquist S. The heat-shock response. Annu Rev Biochem,1986,55:1151-1191
85. Liu H, Charng Y. Acquired thermotolerance independent of heat shock factor Al (HsfAl), the master regulator of the heat stress response. Plant Signal Behav,2012, 7:547-550
86. Liu HC, Charng YY. Common and distinct functions of Arabidopsis class A1 and A2 heat shock factors in diverse abiotic stress responses and development. Plant Physiol, 2013,163:276-290
87. Liu HC, Liao HT, Charng YY. The role of class A1 heat shock factors (HSFA1s) in response to heat and other stresses in Arabidopsis. Plant Cell Environ,2011,34: 738-751
88. Liu HT, Gao F, Li GL, Han JL, De-Long L, Sun DY, Zhou RG. The calmodulin-binding protein kinase 3 is part of heat-shock signal transduction in Arabidopsis thaliana. Plant J,2008,55:760-773
89. Liu HT, Li GL, Chang H, Sun DY, Zhou RG, Li B. Calmodulin-binding protein phosphatase PP7 is involved in thermotolerance in Arabidopsis. Plant Cell Environ, 2007,30:156-164
90. Liu HT, Sun DY, Zhou RG. Ca2+ and AtCaM3 are involved in the expression of heat shock protein gene in Arabidopsis. Plant Cell Environ,2005,28:1276-1284
91. Liu J, Sun N, Liu M, Liu J, Du B, Wang X, Qi X. An autoregulatory loop controlling Arabidopsis HsfA2 expression: role of heat shock-induced alternative splicing. Plant Physiol,2013,162:512-521
92. Liu JG, Qin QL, Zhang Z, Peng RH, Xiong AS, Chen JM, Yao QH. OsHSF7 gene in rice, Oryza sativa L., encodes a transcription factor that functions as a high temperature receptive and responsive factor. BMB Rep,2009,42:16-21
93. Lohmann C, Eggers-Schumacher G, Wunderlich M, Schoffl F. Two different heat shock transcription factors regulate immediate early expression of stress genes in Arabidopsis. Mol Genet Genomics,2004,271:11-21
94. Lomascolo A, Uzan-Boukhris E, Sigoillot JC, Fine F. Rapeseed and sunflower meal: a review on biotechnology status and challenges. Appl Microbiol Biotechnol, 2012, 95:1105-1114
95. Martin T, Schmidt R, Altmann T, Frommer WB. Non-destructive assay systems for detection of β-glucuronidase activity in higher plants. Plant Mol Biol Reporter,1992, 10:37-46
96. Maruyama-Nakashita A, Nakamura Y, Tohge T, Saito K, Takahashi H. Arabidopsis SLIM1 is a central transcriptional regulator of plant sulfur response and metabolism. Plant Cell,2006,18:3235-3251
97. Meiri D, Tazat K, Cohen-Peer R, Farchi-Pisanty O, Aviezer-Hagai K, Avni A, Breiman A. Involvement of Arabidopsis ROF2 (FKBP65) in thermotolerance. Plant Mol Biol,2010,72:191-203
98. Miernyk JA. The J-domain proteins of Arabidopsis thaliana:an unexpectedly large and diverse family of chaperones. Cell Stress Chaperones,2001,6:209-218
99. Mishra SK, Tripp J, Winkelhaus S, Tschiersch B, Theres K, Nover L, Scharf KD. In the complex family of heat stress transcription factors, HSfA1 has a unique role as master regulator of thermotolerance in tomato. Gene Dev,2002,16:1555-1567
100. Mittler R, Finka A, Goloubinoff P. How do plants feel the heat? Trends Biochem Sci, 2012,37:118-125
101. Morrison M. Heat stress during reproduction in summer rape. Can J Bot,1993,71: 303-308
102. Morrison MJ, Stewart DW. Heat stress during flowering in summer Brassica. Crop Sci,2002,42:797-803
103. Mun JH, Kwon SJ, Yang TJ, Seol YJ, Jin M, Kim JA, Lim MH, Kim JS, Baek S, Choi BS. Genome-wide comparative analysis of the Brassica rapa gene space reveals genome shrinkage and differential loss of duplicated genes after whole genome triplication. Genome Biol,2009,10:R1ll
104. Murphy DJ, Cummins I. Biosynthesis of seed storage products during embryogenesis in rapeseed, Brassica napus. J Plant Physiol,1989,135:63-69
105.Nemhauser JL, Feldman LJ, Zambryski PC. Auxin and ETTIN in Arabidopsis gynoecium morphogenesis. Development,2000,127:3877-3888
106. Nesi N, Lucas MO, Auger B, Baron C, Lecureuil A, Guerche P, Kronenberger J, Lepiniec L, Debeaujon I, Renard M. The promoter of the Arabidopsis thaliana BAN gene is active in proanthocyanidin-accumulating cells of the Brassica napus seed coat. Plant Cell Rep,2009,28:601-617
107. Nishizawa A, Yabuta Y, Yoshida E, Maruta T, Yoshimura K, Shigeoka S. Arabidopsis heat shock transcription factor A2 as a key regulator in response to several types of environmental stress. Plant J,2006,48:535-547
108. Nishizawa-Yokoi A, Nosaka R, Hayashi H, Tainaka H, Maruta T, Tamoi M, Ikeda M, Ohme-Takagi M, Yoshimura K, Yabuta Y, Shigeoka S. HsfAld and HsfAle involved in the transcriptional regulation of HsfA2 function as key regulators for the Hsf signaling network in response to environmental stress. Plant Cell Physiol,2011,52: 933-945
109. Nishizawa-Yokoi A, Yoshida E, Yabuta Y, Shigeoka S. Analysis of the regulation of target genes by an Arabidopsis heat shock transcription factor, HsfA2. Biosci Biotechnol Biochem,2009,73:890-895
110. Noh YS, Amasino RM. Regulation of developmental senescence is conserved between Arabidopsis and Brassica napus. Plant Mol Biol,1999,41:195-206
111. Nour-Eldin HH, Andersen TG, Burow M, Madsen SR, J?rgensen ME, Olsen CE, Dreyer I, Hedrich R, Geiger D, Halkier BA. NRT/PTR transporters are essential for translocation of glucosinolate defence compounds to seeds. Nature,2012,488: 531-534
112.Nover L, Bharti K, Doering P, Mishra SK, Ganguli A, Scharf KD. Arabidopsis and the heat stress transcription factor world: How many heat stress transcription factors do we need? Cell Stress Chaperones,2001,6:177-189
113. Panikulangara TJ, Eggers-Schumacher G, Wunderlich M, Stransky H, Schoffl F. Galactinol synthasel. A novel heat shock factor target gene responsible for heat-induced synthesis of raffinose family oligosaccharides in Arabidopsis. Plant Physiol,2004,136:3148-3158
114. Parcy F, Valon C, Raynal M, Gaubier-Comella P, Delseny M, Giraudat J. Regulation of gene expression programs during Arabidopsis seed development:roles of the ABI3 locus and of endogenous abscisic acid. Plant Cell,1994,6:1567-1582
115. Park CH, Kim TW, Son SH, Hwang JY, Lee SC, Chang SC, Kim SH, Kim SW, Kim SK. Brassinosteroids control AtEXPA5 gene expression in Arabidopsis thaliana. Phytochemistry,2010,71:380-387
116. Park HY, Seok HY, Woo DH, Lee SY, Tarte VN, Lee EH, Lee CH, Moon YH. AtERF71/HRE2 transcription factor mediates osmotic stress response as well as hypoxia response in Arabidopsis. Biochem Biophys Res Commun,2011,414: 135-141
117. Park JH, Lee SY, Kim WY, Jung YJ, Chae HB, Jung HS, Kang CH, Shin MR, Kim SY, Su'udi M, Yun DJ, Lee KO, Kim MG, Lee SY. Heat-induced chaperone activity of serine/threonine protein phosphatase 5 enhances thermotolerance in Arabidopsis thaliana. New Phytol,2011,191:692-705
118. Paulsen GM. High temperature responses of crop plants. In: Boote KJ, Physiology and determination of crop yield. University of Minnesota: American Society of Agronomy,1994.365-389
119. Peet M, Sato S, Gardner R. Comparing heat stress effects on male-fertile and male-sterile tomatoes. Plant Cell Environ,1998,21:225-231
120. Peltonen-Sainio P, Jauhiainen L, Trnka M, Olesen JE, Calanca P, Eckersten H, Eitzinger J, Gobin A, Kersebaum KC, Kozyra J. Coincidence of variation in yield and climate in Europe. Agric Ecosyst Environ,2010, 139:483-489
121. Perez DE, Hoyer JS, Johnson AI, Moody ZR, Lopez J, Kaplinsky NJ. BOBBER1 is a noncanonical Arabidopsis small heat shock protein required for both development and thermotolerance. Plant Physiol,2009,151:241-252
122. Poschet G, Hannich B, Raab S, Jungkunz I, Klemens PA, Krueger S, Wic S, Neuhaus HE, Biittner M. A novel Arabidopsis vacuolar glucose exporter is involved in cellular sugar homeostasis and affects the composition of seed storage compounds. Plant Physiol,2011,157:1664-1676
123. Prandl R, Hinderhofer K, Eggers-Schumacher G, Schoffl F. HSF3, a new heat shock factor from Arabidopsis thaliana, derepresses the heat shock response and confers thermotolerance when overexpressed in transgenic plants. Mol Gen Genet, 1998,258: 269-278
124. Prasad BD, Goel S, Krishna P. In silico identification of carboxylate clamp type tetratricopeptide repeat proteins in Arabidopsis and rice as putative co-chaperones of Hsp90/Hsp70. PLoS ONE, 2010,5:e12761
125. Qu C, Fu F, Lu K, Zhang K, Wang R, Xu X, Wang M, Lu J, Wan H, Zhanglin T. Differential accumulation of phenolic compounds and expression of related genes in black- and yellow-seeded Brassica napus. J Exp Bot,2013,64:2885-2898
126. Queitsch C, Hong SW, Vierling E, Lindquist S. Heat shock protein 101 plays a crucial role in thermotolerance in Arabidopsis. Plant Cell,2000,12:479-492
127. Rasbery JM, Shan H, LeClair RJ, Norman M, Matsuda SP, Bartel B. Arabidopsis thaliana squalene epoxidase 1 is essential for root and seed development. JBiol Chem,2007,282:17002-17013
128.Rivero RM, Ruiz JM, Garcia PC, Lopez-Lefebre LR, Sanchez E, Romero L. Resistance to cold and heat stress:accumulation of phenolic compounds in tomato and watermelon plants. Plant Sci, 2001,160:315-321
129. Rizhsky L, Liang H, Mittler R. The combined effect of drought stress and heat shock on gene expression in tobacco. Plant Physiol,2002,130:1143-1151
130. Rizhsky L, Liang H, Shuman J, Shulaev V, Davletova S, Mittler R. When defense pathways collide. The response of Arabidopsis to a combination of drought and heat stress. Plant Physiol,2004,134:1683-1696
131. Romero HM, Berlett BS, Jensen PJ, Pell EJ, Tien M. Investigations into the role of the plastidial peptide methionine sulfoxide reductase in response to oxidative stress in Arabidopsis. Plant Physiol,2004,136:3784-3794
132. Rose J, Catala C, Urbanowicz B. Carbohydrate binding plant hydrolases which alter plant cell walls. USA Patent,0333223 Al.2010-12-30
133. Rowland O, Zheng H, Hepworth SR, Lam P, Jetter R, Kunst L. CER4 encodes an alcohol-forming fatty acyl-coenzyme A reductase involved in cuticular wax production in Arabidopsis. Plant Physiol,2006,142:866-877
134. Saini H, Sedgley M, Aspinall D. Effect of heat stress during floral development on pollen tube growth and ovary anatomy in wheat (Triticum aestivum L.). Funct Plant Biol,1983,10:137-144
135. Sakuma Y, Maruyama K, Osakabe Y, Qin F, Seki M, Shinozaki K, Yamaguchi-Shinozaki K. Functional analysis of an Arabidopsis transcription factor, DREB2A, involved in drought-responsive gene expression. Plant Cell,2006a, 18: 1292-1309
136. Sakuma Y, Maruyama K, Qin F, Osakabe Y, Shinozaki K, Yamaguchi-Shinozaki K. Dual function of an Arabidopsis transcription factor DREB2A in water-stress-responsive and heat-stress-responsive gene expression. Proc Natl Acad Sci USA,2006b,103:18822-18827
137. Samsel KA. The role of chloroplast-localized HSP21 in the stress responses of Arabidopsis thaliana. [Bachelors degree]. The University of Arizona: 2010
138. Sato S, Peet MM, Thomas JF. Determining critical pre- and post-anthesis periods and physiological processes in Lycopersicon esculentum Mill, exposed to moderately elevated temperatures. JExp Bot,2002,53:1187-1195
139. Scharf KD, Berberich T, Ebersberger I, Nover L. The plant heat stress transcription factor (Hsf) family: structure, function and evolution. Biochim Biophys Acta,2011, 1819:104-119
140. Scharf KD, Heider H, Hohfeld I, Lyck R, Schmidt E, Nover L. The tomato Hsf system: HsfA2 needs interaction with HsfAl for efficient nuclear import and may be localized in cytoplasmic heat stress granules. Mol Cell Biol, 1998,18:2240-2251
141. Scharf KD, Siddique M, Vierling E. The expanding family of Arabidopsis thaliana small heat stress proteins and a new family of proteins containing α-crystallin domains (Acd proteins). Cell Stress Chaperones,2001,6:225-237
142.Schlesinger MJ. Heat shock proteins. JBiol Chem,1990,265:12111-12114
143. Schramm F, Ganguli A, Kiehlmann E, Englich G, Walch D, von Koskull-Doring P. The heat stress transcription factor HsfA2 serves as a regulatory amplifier of a subset of genes in the heat stress response in Arabidopsis. Plant Mol Biol,2006,60: 759-772
144. Schramm F, Larkindale J, Kiehlmann E, Ganguli A, Englich G, Vierling E, von Koskull-Doering P. A cascade of transcription factor DREB2A and heat stress transcription factor HsfA3 regulates the heat stress response of Arabidopsis. Plant J, 2008,53:264-274
145. Shonnard GC, Gepts P. Genetics of heat tolerance during reproductive development in common bean. Crop Sci,1994,34:1168-1175
146. Siddique M, Gernhard S, von Koskull-Doering P, Vierling E, Scharf KD. The plant sHSP superfamily: five new members in Arabidopsis thaliana with unexpected properties. Cell Stress Chaperones,2008,13:183-197
147. S?nderby IE, Geu-Flores F, Halkier BA. Biosynthesis of glucosinolates-gene discovery and beyond. Trends Plant Sci,2010,15:283-290
148. Su PH, Li HM. Arabidopsis stromal 70-kD heat shock proteins are essential for plant development and important for thermotolerance of germinating seeds. Plant Physiol, 2008,146:1231-1241
149. Subramanian A, Tamayo P, Mootha VK, Mukherjee S, Ebert BL, Gillette MA, Paulovich A, Pomeroy SL, Golub TR, Lander ES. Gene set enrichment analysis:a knowledge-based approach for interpreting genome-wide expression profiles. Proc Natl Acad Sci USA,2005,102:15545-15550
150. Sugio A, Dreos R, Aparicio F, Maule AJ. The cytosolic protein response as a subcomponent of the wider heat shock response in Arabidopsis. Plant Cell,2009,21: 642-654
151. Suzuki N, Bajad S, Shuman J, Shulaev V, Mittler R. The transcriptional co-activator MBFlc is a key regulator of thermo tolerance in Arabidopsis thaliana. J Biol Chem, 2008,283:9269-9275
152. Suzuki N, Rizhsky L, Liang HJ, Shuman J, Shulaev V, Mittler R. Enhanced tolerance to environmental stress in transgenic plants expressing the transcriptional coactivator multiprotein bridging factor 1c. Plant Physiol,2005,139:1313-1322
153. Suzuki N, Sejima H, Tam R, Schlauch K, Mittler R. Identification of the MBF1 heat-response regulon of Arabidopsis thaliana. Plant J,2011,66:844-851
154. Swindell WR, Huebner M, Weber AP. Transcriptional profiling of Arabidopsis heat shock proteins and transcription factors reveals extensive overlap between heat and non-heat stress response pathways. BMC Genomics,2007,8:125
155. Tanabe N, Yoshimura K, Kimura A, Yabuta Y, Shigeoka S. Differential expression of alternatively spliced mRNAs of Arabidopsis SR protein homologs, atSR30 and atSR45a, in response to environmental stress. Plant Cell Physiol,2007,48: 1036-1049
156. Tayo T, Morgan D. Quantitative analysis of the growth, development and distribution of flowers and pods in oilseed rape (Brassica napus L.). JAgric Sci, Cambridge,1975,85:103-110
157. Thomas J, Palusa SG, Prasad KV, Ali GS, Surabhi GK, Ben-Hur A, Abdel-Ghany SE, Reddy AS. Identification of an intronic splicing regulatory element involved in auto-regulation of alternative splicing of SCL33 pre-mRNA. Plant J,2012,72: 935-946
158. Trenberth KE, Jones PD., Ambenje P, Bojariu R, Easterling D, Tank AK, Parker D, Rahimzadeh F, Renwick JA, Rusticucci M, Soden B and Zhai P. Observations: surface and atmospheric climate change. In: Solomon, S., Qin, D., Manning, M., Chen, Z., Marquis, M., Averyt, K.B., Tignor, M.,Miller, H.L. eds., Climate change 2007:the physical science basis. Contribution of working group I to the fourth assessment report of the Intergovernmental Panel on Climate Change. Cambridge, United Kingdom and New York, NY, USA: Cambridge University Press,2007.
159. Trick M, Cheung F, Drou N, Fraser F, Lobenhofer EK, Hurban P, Magusin A, Town CD, Bancroft I. A newly-developed community microarray resource for transcriptome profiling in Brassica species enables the confirmation of Brassica-specific expressed sequences. BMC Plant Biol,2009,9:50
160. Tsutsui T, Kato W, Asada Y, Sako K, Sato T, Sonoda Y, Kidokoro S, Yamaguchi-Shinozaki K, Tamaoki M, Arakawa K. DEAR1, a transcriptional repressor of DREB protein that mediates plant defense and freezing stress responses in Arabidopsis. J Plant Res,2009,122:633-643
161. Vivian-Smith A, Koltunow AM. Genetic analysis of growth-regulator-induced parthenocarpy in Arabidopsis. Plant Physiol,1999,121:437-452
162. Vogel JT, Zarka DG, Van Buskirk HA, Fowler SG, Thomashow MF. Roles of the CBF2 and ZAT12 transcription factors in configuring the low temperature transcriptome of Arabidopsis. Plant J,2005,41:195-211
163. Volkov RA, Panchuk Ⅱ, Mullineaux PM, Schoeffl F. Heat stress-induced H2O2 is required for effective expression of heat shock genes in Arabidopsis. Plant Mol Biol, 2006,61:733-746
164. Wang X, Xu L, Zheng L. Cloning and expression of phosphatidylcholine-hydrolyzing phospholipase D from Ricinus communis L. J Biol Chem,1994,269:20312-20317
165. Watanabe N, Lam E. Arabidopsis Bax inhibitor-1 functions as an attenuator of biotic and abiotic types of cell death. Plant J,2006,45:884-894
166. Whitecross M, Armstrong D. Environmental effects on epicuticular waxes of Brassica napus L. Aust J Bot,1972,20:87-95
167. Wingenter K, Schulz A, Wormit A, Wic S, Trentmann O, Hoermiller II, Heyer AG, Marten I, Hedrich R, Neuhaus HE. Increased activity of the vacuolar monosaccharide transporter TMT1 alters cellular sugar partitioning, sugar signaling, and seed yield in Arabidopsis. Plant Physiol,2010,154:665-677
168. Wu C. Heat shock transcription factors:structure and regulation. Annu Rev Cell Dev Biol,1995,11:441-469
169. Wu CC, Singh P, Chen MC, Zimmerli L. L-Glutamine inhibits beta-aminobutyric acid-induced stress resistance and priming in Arabidopsis. JExp Bot,2010,61: 995-1002
170. Wu SJ, Wang LC, Yeh CH, Lu CA, Wu SJ. Isolation and characterization of the Arabidopsis heat-intolerant 2 (hit2) mutant reveal the essential role of the nuclear export receptor EXPORTTN1A (XPO1A) in plant heat tolerance. New Phytol,2010, 186:833-842
171. Wu TY, Juan YT, Hsu YH, Wu SH, Liao HT, Fung RW, Charng YY. Interplay between heat shock proteins, HSP101 and HSA32, prolongs heat acclimation memory posttranscriptionally in Arabidopsis. Plant Physiol,2013,161:2075-2084
172. Wunderlich M, Werr W, Schoffl F. Generation of dominant-negative effects on the heat shock response in Arabidopsis thaliana by transgenic expression of a chimaeric HSF1 protein fusion construct. Plant J,2003,35:442-451
173. Xin H, Zhang H, Chen L, Li X, Lian Q, Yuan X, Hu X, Cao L, He X, Yi M. Cloning and characterization of HsfA2 from Lily (Lilium longiflorum). Plant Cell Rep,2010, 29:875-885
174. Xing Y, Jia W, Zhang J. AtMKK1 and AtMPK6 are involved in abscisic acid and sugar signaling in Arabidopsis seed germination. Plant Mol Biol,2009,70:725-736
175. Xu X, Song H, Zhou Z, Shi N, Ying Q, Wang H. Functional characterization of AtHsp90.3 in Saccharomyces cerevisiae and Arabidopsis thaliana under heat stress. Biotechnol Lett,2010,32:979-987
176. Xuan Y, Zhou S, Wang L, Cheng Y, Zhao L. Nitric oxide functions as a signal and acts upstream of AtCaM3 in thermotolerance in Arabidopsis seedlings. Plant Physiol, 2010,153:1895-1906
177. Yamada K, Fukao Y, Hayashi M, Fukazawa M, Suzuki I, Nishimura M. Cytosolic HSP90 regulates the heat shock response that is responsible for heat acclimation in Arabidopsis thaliana. J Biol Chem,2007,282:37794-37804
178. Yamakawa H, Hakata M. Atlas of rice grain filling-related metabolism under high temperature:joint analysis of metabolome and transcriptome demonstrated inhibition of starch accumulation and induction of amino acid accumulation. Plant Cell Physiol, 2010,51:795-809
179. Yamakawa H, Hirose T, Kuroda M, Yamaguchi T. Comprehensive expression profiling of rice grain filling-related genes under high temperature using DNA microarray. Plant Physiol,2007,144:258-277
180. Yamamoto Y, Shimada Y, Kimura M, Manabe K, Sekine Y, Matsui M, Ryuto H, Fukunishi N, Abe T, Yoshida S. Global classification of transcriptional responses to light stress in Arabidopsis thaliana. Endocytobiosis Cell Res,2004,15:438-452
181. Yang C Y, Hsu FC, Li JP, Wang NN, Shih MC. The AP2/ERF transcription factor AtERF73/HRE1 modulates ethylene responses during hypoxia in Arabidopsis. Plant Physiol,2011,156:202-212
182. Yang K, Lim CJ, Hong JK, Park CY, Cheong YH, Chung WS, Lee KO, Lee S Y, Cho MJ, Lim CO. Identification of cell wall genes modified by a permissive high temperature in Chinese cabbage. Plant Sci,2006,171:175-182
183. Yang YW, Lai KN, Tai PY, Li WH. Rates of nucleotide substitution in angiosperm mitochondrial DNA sequences and dates of divergence between Brassica and other angiosperm lineages. J Mol Evol,1999,48:597-604
184. Yokotani N, Ichikawa T, Kondou Y, Matsui M, Hirochika H, Iwabuchi M, Oda K. Expression of rice heat stress transcription factor OsHsfA2e enhances tolerance to environmental stresses in transgenic Arabidopsis. Planta,2008,227:957-967
185. Yoo CY, Miura K, Jin JB, Lee J, Park HC, Salt DE, Yun DJ, Bressan RA, Hasegawa PM. SIZ1 small ubiquitin-like modifier E3 ligase facilitates basal thermotolerance in Arabidopsis independent of salicylic acid. Plant Physiol,2006,142:1548-1558
186. Yoo SD, Cho YH, Sheen J. Arabidopsis mesophyll protoplasts:a versatile cell system for transient gene expression analysis. Nat Protoc,2007,2:1565-1572
187. Yoshida T, Sakuma Y, Todaka D, Maruyama K, Qin F, Mizoi J, Kidokoro S, Fujita Y, Shinozaki K, Yamaguchi-Shinozaki K. Functional analysis of an Arabidopsis heat-shock transcription factor HsfA3 in the transcriptional cascade downstream of the DREB2A stress-regulatory system. Biochem Biophys Res Commun,2008,368: 515-521
188. Yoshimura K, Mori T, Yokoyama K, Koike Y, Tanabe N, Sato N, Takahashi H, Maruta T, Shigeoka S. Identification of alternative splicing events regulated by an Arabidopsis serine/arginine-like protein, atSR45a, in response to high-light stress using a tiling array. Plant Cell Physiol,2011,52:1786-1805
189. Young LW, Wilen RW, Bonham-Smith PC. High temperature stress of Brassica napus during flowering reduces micro- and megagametophyte fertility, induces fruit abortion, and disrupts seed production. JExp Bot,2004,55:485-495
190. Yu HD, Yang XF, Chen ST, Wang YT, Li JK, Shen Q, Liu XL, Guo FQ. Downregulation of chloroplast RPS1 negatively modulates nuclear heat-responsive expression of HsfA2 and its target genes in Arabidopsis. Plos Genet,2012a,8: e1002669
191. Yu X, Wang H, Lu Y, de Ruiter M, Cariaso M, Prins M, van Tunen A, He Y. Identification of conserved and novel microRNAs that are responsive to heat stress in Brassica rapa. J Exp Bot, 2012b,63:1025-1038
192. Zanewich KP, Rood SB. Distribution of endogenous gibberellins in vegetative and reproductive organs of Brassica. J Plant Growth Regul,1993,12:41-46
193. Zelisko A, Garcia-Lorenzo M, Jackowski G, Jansson S, Funk C. AtFtsH6 is involved in the degradation of the light-harvesting complex II during high-light acclimation and senescence. Proc Natl Acad Sci USA,2005,102:13699-13704
194. Zhang H, Yang J, Chen Y, Mao X, Wang Z, Li C. Identification and expression analysis of the heat shock transcription factor (HSF) gene family in Populus trichocarpa. Plant Omics.2013,6:415-424
195. Zhang LR, Li YS, Xing D, Gao CJ. Characterization of mitochondrial dynamics and subcellular localization of ROS reveal that HsfA2 alleviates oxidative damage caused by heat stress in Arabidopsis. J Exp Bot,2009a,60:2073-2091
196. Zhang W, Zhou RG, Gao YJ, Zheng SZ, Xu P, Zhang SQ, Sun DY. Molecular and genetic evidence for the key role of AtCaM3 in heat-shock signal transduction in Arabidopsis. Plant Physiol,2009b,149:1773-1784
197. Zhou S, Zhang P, Jing Z, Shi J. Genome-wide identification and analysis of heat shock transcription factor family in cucumber (Cucumis sativus L.). Plant Omics. 2013,6:449-455
198. Zhou W, Zhou T, Li MX, Zhao CL, Jia N, Wang XX, Sun YZ, Li GL, Xu M, Zhou RG. The Arabidopsis J-protein AtDjBl facilitates thermotolerance by protecting cells against heat-induced oxidative damage. New Phytol, 2012,194:364-378
199. Zhu G, Liu Y, Ye N, Liu R, Zhang J. Involvement of the abscisic acid catabolic gene CYP707A2 in the glucose-induced delay in seed germination and post-germination growth of Arabidopsis. Physiol Plant,2011,143:375-384
200. Zhu Y, Cao Z, Xu F, Huang Y, Chen M, Guo W, Zhou W, Zhu J, Meng J, Zou J. Analysis of gene expression profiles of two near-isogenic lines differing at a QTL region affecting oil content at high temperatures during seed maturation in oilseed rape(Brassica napus L.). Theor Appl Genet,2012,124:515-531
201. Zhu Y, Wang Z, Jing Y, Wang LL, Liu X, Liu YX, Deng X. Ectopic over-expression of BhHsfl, a heat shock factor from the resurrection plant Boea hygrometrica, leads to increased thermotolerance and retarded growth in transgenic Arabidopsis and tobacco. Plant Mol Biol,2009a,71:451-467
202. Zimmerli L, Hou BH, Tsai CH, Jakab G, Mauch-Mani B, Somerville S. The xenobiotic beta-aminobutyric acid enhances Arabidopsis thermotolerance. Plant J, 2008,53:144-156