水稻芽期响应镉胁迫的转录组分析

doi:10.7668/hbnxb.20193136

摘要/Abstract

摘要：

为研究不同类型水稻对镉(Cd)胁迫差异的原因,以镉敏感籼稻恢复系昌恢891(CH891)和镉耐受粳稻品种02428为材料,选取种子萌发后,连续镉处理3 d的和未进行镉处理的幼芽,使用高通量Illumina HiSeq 4000测序技术进行RNA-Seq分析,共得到539 524 490个有效读长,与参考基因组的比对率在94.81%~96.82%,GC含量均在49%以上。通过比较分析,共检测到7 204个差异表达基因(DEGs),其中在CH891中特异表达的基因(SCR3)有849个,02428中特异表达的基因(RCR3)有676个,2个品种均有表达但表达量存在差异的基因(CCR3)有770个。KEGG和GO富集通路分析表明,异黄酮生物合成、磷酸肌醇代谢、黄酮类生物合成和花青素生物合成途径在SCR3中显著富集,细胞核、自噬调节和胰岛素抗性途径在RCR3中显著富集,α-亚麻酸代谢、吞噬体、柠檬烯和蒎烯降解和脂肪酸降解途径在CCR3中显著富集。结果表明,Cd处理后,次级代谢过程主要在CH891中富集,蛋白质代谢主要在02428中富集,综上所述,CH891和02428响应的代谢路径存在差异,控制这些代谢路径的分子机制可能是造成不同水稻镉胁迫差异的主要原因。

关键词: 籼稻, 粳稻, 镉胁迫, 转录组, 差异表达

Abstract:

In order to study the reasons for the differences of different types of rice under cadmium stress,it used the cadmium sensitive indica rice restorer line Changhui 891(CH891)and the cadmium tolerant japonica rice variety 02428 as materials,and selected the buds after seed germination,which were continuously treated with cadmium for 3 days and not treated with cadmium for RNA-Seq analysis using high-throughput Illumina HiSeq 4000 sequencing technology.A total of 539 524 490 valid read were obtained,the comparison rate with the reference genome was between 94.81% and 96.82%,and the GC content was above 49%.Through comparative analysis,a total of 7 204 differentially expressed genes(DEGs)were detected,among which 849 genes were specifically expressed in CH891(SCR3),676 genes were specifically expressed in 02428(RCR3),and 770 genes were expressed in both varieties(CCR3),but the expression level was different.The analysis of enrichment pathways of KEGG and GO showed that isoflavone biosynthesis,inositol phosphate metabolism,flavonoid biosynthesis and anthocyanin biosynthesis were significantly enriched in SCR3.Ribosome,regulation of autophagy and insulin resistance pathways were significantly enriched in RCR3.Alpha-linolenic acid metabolism,phagosome,limonene and pinene degradation and fatty acid degradation pathways were significantly enriched in CCR3.The results showed that after Cd treatment,the secondary metabolic process was mainly enriched in CH891,and the protein metabolism was mainly enriched in 02428.In conclusion,the metabolic pathways of CH891 and 02428 were different after Cd treatment,and the molecular mechanism controlling these metabolic pathways might be the main reason for the differences of cadmium stress in different rice varieties.

Key words: Indica rice, Japonica rice, Cadmium stress, Transcriptome, Differential expression

王燕宁, 黄涛, 吴光亮, 黄诗颖, 钟奇, 王鹏, 杨朦朦, 李才敬, 程琴, 贺浩华, 金伟, 郭凌, 边建民. 水稻芽期响应镉胁迫的转录组分析[J]. 华北农学报, 2022, 37(5): 25-35. doi: 10.7668/hbnxb.20193136.

WANG Yanning, HUANG Tao, WU Guangliang, HUANG Shiying, ZHONG Qi, WANG Peng, YANG Mengmeng, LI Caijing, CHENG Qin, HE Haohua, JIN Wei, GUO Ling, BIAN Jianmin. Transcriptome Analysis of Rice Bud Stage in Response to Cadmium Stress[J]. Acta Agriculturae Boreali-Sinica, 2022, 37(5): 25-35. doi: 10.7668/hbnxb.20193136.

http://www.hbnxb.net/CN/Y2022/V37/I5/25

图/表 13

表1 qRT-PCR引物序列

Tab.1 The sequences for qRT-PCR primers

基因名称 Gene name	正向引物(5'—3') F-primer(5'—3')	反向引物(5'—3') R-primer(5'—3')
LOC_Os03g08470	CTGGAGTTCGACCCGTTCAT	TCGAAGCTCCACAGGTTGAC
LOC_Os04g39880	CACCTCAGGCGTTGGAAGAT	AGGCCAGTTGCATAGCCATT
LOC_Os09g36700	TGCCATTGCTGCCAAAGATG	GGACAGGGAAGAGTTGGTGG
LOC_Os10g40720	CAGGTGGATCTCATGGAGGC	GAAAGAACGGTAGACGGCCA
LOC_Os02g02400	GGACGAGGAGGTGGACTACT	TGCTTGTGTATCGTCGCCTT
LOC_Os04g33830	TTTCCGCAGGACTGGCTTAG	GAGAAGAAGAGCCCCGTGAG
LOC_Os06g21590	ATGGAGAAGGACCCCGAGAA	ACGCAGAAACCCACAAAAGC
LOC_Os09g31430	TGAGGGCGCCTACAAAGAAG	GGAGAACCGGAAAGCATCCA
LOC_Os01g18744	CGATTCCCTGTTCGTCTCCG	TGAACGGGTAGTACGCGAAG
LOC_Os06g13280	GTTGGGGAGAGGAGGATTGC	CAGACCCGACCACCGTATTT
LOC_Os03g03600	CAAGCAGATGCCGTACAAGC	TCCTCGAACGTCTTGAGCAC
LOC_Os08g06110	GAGGGTCGTCTGGCTTTTGA	ACCGTCACCGTGGTTACTTC

图1 CH891和02428在不同处理条件下的表现 A.对照处理;B.镉处理;C.苗高;不同小写字母表示处理组与对照组相比苗高差异显著(P<0.05)。

Fig.1 CH891 and 02428 performance under different treatment conditions A.Control treatment;B.Cadmium treatment;C.Seedling height;Different lowercase letters indicate that the seedling height of the treatment group is significantly different from that of the control group(P<0.05).

表2 测序数据统计和参考基因组上的比对率

Tab.2 Sequencing data statistics and mapped ratios to the reference genome

样本 Sample	原始测序读长 Raw read	原始测序数据/G Raw base	有效测序读长 Vaild read	有效数据/G Valid base	有效读长占比/% Valid ratio	Q20/%	Q30/%	GC含量/% GC content	匹配读长 Mapped reads	唯一匹配读长 Unique Mapped reads
CK_02428_A	45 696 550	6.85	45 295 626	6.79	99.12	99.75	97.20	51.0	43 662 999(96.40%)	29 723 198(65.62%)
CK_02428_B	43 930 280	6.59	43 488 498	6.52	98.99	99.73	97.14	49.0	41 938 316(96.44%)	32 091 614(73.79%)
CK_02428_C	47 475 320	7.12	47 064 332	7.06	99.13	99.71	97.09	50.0	45 444 073(96.56%)	34 732 722(73.80%)
Cd_02428_A	45 814 010	6.87	45 396 612	6.81	99.09	99.62	96.82	50.0	43 807 260(96.50%)	33 435 746(73.65%)
Cd_02428_B	38 458 390	5.77	38 150 196	5.72	99.20	99.74	97.21	50.0	36 936 246(96.82%)	28 311 301(74.21%)
Cd_02428_C	43 063 576	6.46	42 676 692	6.40	99.10	99.70	96.94	49.5	41 266 571(96.70%)	31 487 488(73.78%)
CK_CH891_A	46 690 704	7.00	46 101 030	6.92	98.74	99.74	97.36	49.5	43 707 017(94.81%)	29 993 285(65.06%)
CK_CH891_B	57 839 194	8.68	56 921 950	8.54	98.41	99.75	97.56	50.0	53 993 658(94.86%)	35 354 036(62.11%)
CK_CH891_C	40 120 618	6.02	39 672 946	5.95	98.88	99.74	97.44	50.0	37 650 255(94.90%)	26 731 113(67.38%)
Cd_CH891_A	50 374 398	7.56	49 926 492	7.49	99.11	99.76	97.37	51.0	47 493 447(95.13%)	34 023 070(68.15%)
Cd_CH891_B	41 353 200	6.20	40 980 030	6.15	99.10	99.73	97.21	50.0	38 855 388(94.82%)	27 810 209(67.86%)
Cd_CH891_C	44 193 960	6.63	43 850 086	6.58	99.22	99.72	97.27	51.0	41 624 578(94.92%)	29 506 145(67.29%)

图2 所有比较组差异表达基因 A、B、C和D.火山分布图;E.上下调差异表达基因个数。

Fig.2 DEGs in all comparison groups A,B,C and D.Volcano plot;E.The number of up and down-regulated DEGs.

图3 差异表达基因转录组与qRT-PCR比较

Fig.3 Comparison of differentially expressed gene transcriptome and qRT-PCR results

图4 4个比较组差异表达基因的韦恩图

Fig.4 Venn diagrams for DEGs in the four comparison groups

表3 差异表达基因的分类

Tab.3 Classification of three categories of DEGs

分类 Categories	亚组 Subgroups	差异表达基因数 Number of DEGs
SCR3	仅Cd3_CH891vs.CK3_CH891、	715
	Cd3_CH891vs.CK3_CH891、Cd3_CH891vs.Cd3_02428	134
RCR3	仅Cd3_02428vs.CK3_02428、	565
	Cd3_02428vs.CK3_02428、Cd3_CH891vs.Cd3_02428	111
CCR3	仅Cd3_CH891vs.Cd3_02428	506
	Cd3_CH891vs.CK3_CH891、Cd3_02428vs.CK3_02428、	253
	Cd3_CH891vs.CK3_CH891、Cd3_02428vs.CK3_02428、	11
	Cd3_CH891vs.Cd3_02428

图5 SCR3中的差异表达基因的GO富集 1.生物进程;2.转录调节、DNA模板;3.转录、DNA模板;4.蛋白磷酸化;5.代谢过程;6.伤害反应;7.氧化还原反应;8.脱落酸反应;9.油菜素类固醇反应;10.脂质转运;11.缺氧反应;12.蛋白质泛素化;13.蛋白酶体介导的泛素化依赖性蛋白质分解代谢过程;14.冷反应;15.多细胞生物发育;16.种子休眠中组织胚胎发育;17.叶发育;18.光刺激反应;19.蛋白质折叠;20.类黄酮葡萄糖醛酸化;21.胞外分泌;22.次级代谢生物合成;23.防御反应;24.赤霉素反应;25.甲基化作用;26.核;27.膜整体结构;28.质膜;29.细胞质;30.细胞外区域;31.胞液;32.叶绿体;33.膜;34.高尔基体;35.线粒体;36.细胞壁;37.植物细胞壁;38.内质网;39.质膜固定组成成分;40.胞间连丝;41.分子功能;42.RNA结合;43.转录活性、序列特异性DNA结合;44.金属离子结合;45.蛋白结合;46.ATP结合;47.羧基酯水解酶;48.蛋白丝氨酸/苏氨酸激酶活性;49.锌离子结合;50.水解酶活性。

Fig.5 The GO enrichment of differentially expressed genes in SCR3 1.Biological process;2.Regulation of transcription,DNA-templated;3.Transcription,DNA-templated;4.Protein phosphorylation;5.Metabolic process;6.Response to wounding;7.Oxidation-reduction reaction;8.Response to abscisic acid;9.Response to brassinosteroid;10.Lipid transport;11.Response to hypoxia;12.Protein ubiquitination;13.Proteasome-mediated ubiquitination-dependent catabolic process;14.Response to cold;15.Multicellular organism development;16.Embryo development ending in seed dormancy;17.Leaf development;18.Response to light stimulation reaction;19.Protein folding;20.Flavonold glucuronidation;21.Exocytosis;22.Secondary metabolic biosynthesis;23.Defense response;24.Response to gibberellin;25.Methylation;26.Nucleus;27.Integral component of membrane;28.Plasma membrane;29.Cytoplasm;30.Extracellular region;31.Cytosol;32.Chloroplast;33.Membrane;34.Golgi apparatus;35.Mitochondrion;36.Cell wall;37.Plant-type cell wall;38.Endoplasmic reticulum;39.Anchored components of plasma membrane;40.Plasmodesmata;41.Molecular function;42.RNA binding;43.Transcription factor activity,sequence-specific DNA binding;44.Metal ion binding;45.Protein binding;46.ATP binding;47.Carboxylic ester hydrolase;48.Protein serine/threonine kinase activity;49.Zinc ion binding;50.Hydrolase activity.

图6 SCR3中的差异表达基因的KEGG富集 1.α亚麻酸代谢;2.萜类骨架生物合成;3.类固醇生物合成;4.淀粉和蔗糖代谢;5.倍半萜和三萜类生物合成;6.自噬调节;7.RNA聚合酶;8.RNA降解;9.嘧啶代谢;10.卟啉和叶绿素代谢;11.植物激素信号转导;12.光合作用;13.磷脂酰肌醇信号系统;14.类苯丙酸生物合成;15.苯丙氨酸、酪氨酸和色氨酸生物合成;16.磷酸戊糖途径;17.N-聚糖生物合成;18.异黄酮生物合成;19.磷酸肌醇代谢;20.糖基磷脂酰肌醇生物合成;21.硫代葡萄糖苷生物合成;22.类黄酮生物合成;23.醚类脂代谢;24.二帖生物合成;25.角质、木栓质和蜡质的生物合成;26.类胡萝卜素;27.光合生物的碳固定;28.生物合成;29.碱基切除;30.花青素生物合成。

Fig.6 The KEGG enrichment of differentially expressed genes in SCR3 1.Alpha-Linolenic acid metabolism;2.Terpenoid backbone biosynthesis;3.Steroid biosynthesis;4.Starch and sucrose metabolism;5.Sesquiterpenoid and triterpenoid biosynthesis;6.Regulation of autophagy;7.RNA polymerase;8.RNA degradation;9.Pyrimidine metabolism;10.Porphyrin and chlorophyll metabolism;11.Plant hormone signal transduction;12.Photosynthesis;13.Phosphatidylinositol signaling system;14.Phenylpropanoid biosynthesis;15.Phenylalanine,tyrosine and tryptophan biosynthesis;16.Pentose phosphate pathway;17.N-Glycan biosynthesis;18.Isoflavonoid biosynthesis;19.Inositol phosphate metabolism;20.Glycosylphosphatidylinositol(GPI)-anchor biosynthesis;21.Glucosinolate biosynthesis;22.Flavonoid biosynthesis;23.Ether lipid metabolism;24.Diterpenoid biosynthesis;25.Cutin,suberine and wax biosynthesis;26.Carotenoids;27.Carbon fixation in photosynthetic organisms;28.Biosynthesis;29.Base excision repair;30.Anthocyanin biosynthesis.

图7 RCR3中的差异表达基因的GO富集 1.生物过程;2.转录调控、DNA模板;3.转录、DNA模板;4.氧化还原反应;5.防御反应;6.蛋白磷酸化;7.细胞壁组织;8.脱落酸反应;9.伤害反应;10.盐胁迫反应;11.蛋白质水解;12.多细胞生物发育;13.一维细胞生长;14.氧化应激反应;15.冷反应;16.独脚金内酯生物合成;17.茉莉酸反应;18.蛋白酶体介导的泛素依赖性蛋白质分解代谢过程;19.信号转导;20.脂质转运;21.类黄酮生物合成过程;22.卡里金反应;23.涉及多维细胞生长的植物型细胞壁修饰;24.蛋白转运体;25.胞吐作用中的囊泡对接;26.核;27.质膜;28.膜整体结构;29.胞外区域;30.细胞质;31.线粒体;32.膜;33.叶绿体;34.细胞溶质;35.胞间连丝;36.细胞壁;37.细胞组分;38.液泡;39.植物细胞壁;40.细胞内有膜的细胞器;41.分子功能;42.转录活性、序列特异性DNA结合;43.蛋白结合;44.DNA结合;45.蛋白丝氨酸/苏氨酸激酶活性;46.ATP结合;47.金属离子结合;48.激酶活性;49.核糖体的结构成分;50.锌离子结合。

Fig.7 The GO enrichment of differentially expressed genes in RCR3 1.Biological process;2.Regulation of transcription,DNA-template;3.Transcription,DNA-template;4.Oxidation-reduction reaction;5.Defense response;6.Protein phosphorylation;7.Cell wall organization;8.Response to abscisic acid;9.Response to wounding;10.Response to salt stress;11.Proteolysis;12.Multicellular organism development;13.Undimensional cell growth;14.Response to oxidative stress;15.Response to cold;16.Strigolactone biosynthetic process;17.Response to jasmonic acid;18.Proteasome-mediated ubiquitination-dependent catabolic process;19.Signal transduction;20.Lipid transport;21.Flavonoid biosynthetic process;22.Response to Karikin;23.Plant-type cell wall modification involved in multidimensional cell growth;24.Protein transport;25.Vesicle docking involved in exocytosis;26.Nucleus;27.Plasma membrane;28.Integral component of membrane;29.Extracellular region ;30.Cytoplasm;31.Mitochondria;32.Membrane;33.Chloroplast;34.Cytosol;35.Plasmodesmata;36.Cell wall;37.Cellular components;38.Vacuole;39.Plant-type cell wall;40.Intracellular membrane-bounded organelle;41.Molecular function;42.Transcription factor activity,sequence-specific DNA binding;43.Protein binding;44.DNA binding;45.Protein serine/Threonine kinase activity;46.ATP binding;47.Metal ion binding;48.Kinase activity;49.Structural constituent of ribosome;50.Zinc ion binding.

图8 RCR3中的差异表达基因的KEGG富集 1.mRNA监控途径;2.β-丙氨酸代谢;3.玉米素生物合成;4.缬氨酸、亮氨酸和异亮氨酸降解;5.囊泡运输中的SNARE相互作用;6.核糖体;7.自噬调节;8.丙酸代谢;9.植物-病原体相互作用;10.植物激素信号转导;11.苯丙酸类生物合成;12.苯丙氨酸、酪氨酸和色氨酸的生物合成;13.磷酸戊糖;14.氧化磷酸化;15.其他类型O-聚糖生物合成;16.其他聚糖降解;17.N聚糖生物合成;18.胰岛素抗性;19.糖苷磷酸肌醇生物合成;20.醣酵解/糖异生;21.果糖和甘露糖;22.二帖生物合成;23.角质、木栓质和蜡质的生物合成;24.TCA循环;25.类胡萝卜素生物合成;26.碳代谢;27.碱基切除修复;28.抗坏血酸盐和醛酸盐;29.精氨酸和脯氨酸代谢;30.花青素生物合成。

Fig.8 The KEGG enrichment of differentially expressed genes in RCR3 1.mRNA surveillance pathway;2.Beta-Alanine metabolism;3.Zeatin biosynthesis;4.Valine,leucine and isoleucine degradation;5.SNARE interactions in vesicular transport;6.Ribosome;7.Regulation of autophagy;8.Propanoate metabolism;9.Plant-pathogen interaction;10.Plant hormone signal transduction;11.Phenylpropanoid biosynthesis;12.Phenylalanine,tyrosine and tryptophan biosynthesis;13.Pentose phosphate pathway;14.Oxidative phosphorylation;15.Other types of O-glycan biosynthesis;16.Other glycan degradation;17.N-glycan biosynthesis;18.Insulin resistance;19.Glycosylphosphatidylinositol(GPI)-anchor biosynthesis;20.Glycolysis/Gluconeogenesis;21.Fructose and mannose metabolism;22.Diterpenoid biosynthesis;23.Cutin,suberine and wax biosynthesis;24.Citrate cycle(TCA cycle);25.Carotenoid biosynthesis;26.Carbon metabolism;27.Base excision repair;28.Ascorbate and aldarate metabolism;29.Arginine and proline metabolism;30.Anthocyanin biosynthesis.

图9 CCR3中的差异表达基因的GO富集 1.转录调节、DNA模板;2.生物进程;3.转录、DNA模板;4.伤害反应;5.蛋白质泛素化;6.氧化还原反应;7.冷反应;8.脱落酸反应;9.水杨酸反应;10.寡肽转运;11.防御反应;12.氧化应激反应;13.脂肪酸生物合成;14.脱水反应;15.参与细胞内蛋白质代谢过程的蛋白质水解;16.光刺激反应;17.一维细胞生长;18.乙炔激活信号转导;19.赤霉酸介导的信号转导;20.次级代谢生物合成;21.盐胁迫反应;22.几丁质反应;23.调控防御反应;24.其他有机物反应;25.木质素生物合成;26.核;27.细胞质;28.膜整体结构;29.质膜;30.膜;31.叶绿体;32.细胞内;33.胞外区域;34.线粒体;35.质膜固定成分;36.细胞溶质;37.高尔基体;38.质体;39.叶绿体基质;40.细胞壁;41.转录因子活性、序列特异性DNA结合;42.分子功能;43.蛋白质结合;44.DNA结合;45.锌离子结合;46.序列特异性DNA结合;47.转运蛋白活性;48.转移酶活性,转移糖酰基;49.蛋白泛素化转移酶活性;50.转录调控区DNA结合。

Fig.9 The GO enrichment of differentially expressed genes in CCR3 1.Regulation of transcription,DNA-template;2.Biological process;3.Transcription,DNA-template;4.Response to wounding;5.Protein ubiquitination;6.Oxidation-reduction reaction;7.Response to cold;8.Response to abscisic acid;9.Response to salicylic acid;10.Oligopeptide transport;11.Defense response;12.Response to oxidative stress;13.Fatty acid biosynthetic process;14.Response water dehydration;15.Protelysis involved in cellular protein catabolic process;16.Response to light stimulus;17.Undimensional cell growth;18.Acetylen-activated signaling pathway;19.Gibberellic acid mediated signaling transduction;20.Secondary metabolic biosynthesis;21.Salt stress reaction;22.Chitin reaction;23.Regulating the defense response;24.Other organic reactions;25.Lignin biosynthetic process;26.Nuclus;27.Cytoplasm;28.Integral component of membrane;29.Plasma membrane;30.Membrane;31.Chloroplast;32.Intracellular;33.Extracellular region;34.Mitochondria;35.Anchored components of plasma membrane;36.Cytosol;37.Golgi apparatus;38.Plastid;39.Chloroplast stroma;40.Cell wall;41.Transcription factor activity,sequence-specific DNA binding;42.Molecular function;43.Protein binding;44.DNA binding;45.Zinc ion binding;46.Sequence-specific DNA binding;47.Transporter activity;48.Transferase activity,transferring glycoayl groups;49.Ubiquitin-protein transferase activity;50.Transcription regulatory region DNA binding.

图10 CCR3中的差异表达基因的KEGG富集 1.α亚麻酸代谢;2.缬氨酸、亮氨酸和异亮氨酸降解;3.锑烯类、二芳基庚烷类和姜酚生物合成;4.植物病原体互作;5.吞噬体;6.过氧化物酶体;7.烟酸盐和烟酰胺代谢;8.N聚糖生物合成;9.赖氨酸降解;10.亚油酸代谢;11.柠檬烯和蒎烯降解;12.异黄酮类生物合成;13.糖磷脂生物合成;14.糖磷脂生物合成-神经节系列;15.糖酵解/糖异生;16.甘氨酸、丝氨酸和苏氨酸代谢;17.甘油磷脂代谢;18.半乳糖代谢;19.脂肪酸延伸;20.脂肪酸降解;21.脂肪酸生物合成;22.二帖生物合成;23.角质、木栓质和蜡质的生物合成;24.TCA循环;25.植物昼夜节律;26.碳代谢;27.油菜素类固醇生物合成;28.精氨酸生物合成;29.氨酰基tRNA生物合成;30.丙氨酸、天冬氨酸和谷氨酸代谢。

Fig.10 The KEGG enrichment of differentially expressed genes in CCR3 1.Alpha-Linolenic acid metabolism;2.Valine,leucine and isoleucine degradation;3.Stilbenoid,diarylheptanoid and gingerol biosynthesis;4.Plant-pathogen interaction;5.Phagosome;6.Peroxisome;7.Nicotinate and nicotinamide metabolism;8.N-Glycan biosynthesis;9.Lysine degradation;10.Linoleic acid metabolism;11.Limonene and pinene degradation;12.Isoflavonoid biosynthesis;13.Glycophospholipid biosynthesis-globo series;14.Glycophospholipid biosynthesis-ganglio series;15.Glycolysis/Gluconeogenesis;16.Glycine,serine and threonine metabolism;17.Glycerophospholipid metabolism;18.Galactose metabolism;19.Fatty acid elongation;20.Fatty acid degradation;21.Fatty acid biosynthesis;22.Diterpenoid biosynthesis;23.Cutin,suberine and wax biosythesis;24.Citrate cycle(TCA cycle);25.Circadian rhythm-plant;26.Carbon metabolism;27.Brassinosteroid biosynthesis;28.Arginine biosynthesis;29.Aminoacyl-tRNA biosynthesis;30.Alanine,aspartate and glutamate metabolism.

参考文献 35

[1]	许肖博, 安鹏虎, 郭天骄, 韩丹, 贾玮, 黄五星. 水稻镉胁迫响应机制及防控措施研究进展[J]. 中国水稻科学, 2021, 35(5):415-426.doi: 10.16819/j.1001-7216.2021.201209. doi: 10.16819/j.1001-7216.2021.201209
	Xu X B, An P H, Guo T J, Han D, Jia W, Huang W X. Research progresses on response mechanisms and control measures of cadmium stress in rice[J]. Chinese Journal of Rice Science, 2021, 35(5):415-426. doi: 10.16819/j.1001-7216.2021.201209
[2]	邹文莉. 水稻苗期镉耐性和籽粒低镉积累基因挖掘[D]. 南昌: 江西农业大学, 2019.doi: 10.27177/d.cnki.gjxnu.2019.000131. doi: 10.27177/d.cnki.gjxnu.2019.000131
	Zou W L. Genetic dissection of Cd tolerance at seedling stage and grain low Cd accumulation in rice(Orazy sativa L.)[D]. Nanchang: Jiangxi Agricultural University, 2019.
[3]	张大为, 杜云燕, 吴金锋, 周定港, 刘丽莉, 刘忠松, 严明理. 镉胁迫对甘蓝型油菜幼苗生长及基因表达的影响[J]. 中国油料作物学报, 2020, 42(4):613-622.doi: 10.19802/j.issn.1007-9084.2020100. doi: 10.19802/j.issn.1007-9084.2020100
	Zhang D W, Du Y Y, Wu J F, Zhou D G, Liu L L, Liu Z S, Yan M L. Effect of cadmium stress on plant growth and gene expression in Brassica napus seedlings[J]. Chinese Journal of Oil Crop Sciences, 2020, 42(4):613-622.
[4]	冷艳. 绿豆幼苗响应镉胁迫的分子机制研究[D]. 兰州: 兰州交通大学, 2021.doi: 10.27205/d.cnki.gltec.2021.000034. doi: 10.27205/d.cnki.gltec.2021.000034
	Leng Y. Molecular mechanism of mung bean seedlings in response to cadmium stress[D]. Lanzhou: Lanzhou Jiaotong University, 2021
[5]	Kumar S, Sharma A. Cadmium toxicity:Effects on human reproduction and fertility[J]. Reviews on Environmental Health, 2019, 34(4):327-338.doi: 10.1515/reveh-2019-0016. doi: 10.1515/reveh-2019-0016 URL
[6]	Yan Y F, Lestari P, Lee K J, Kim M Y, Lee S H, Lee B W. Identification of quantitative trait loci for cadmium accumulation and distribution in rice(Oryza sativa)[J]. Genome, 2013, 56(4):227-232.doi: 10.1139/gen-2012-0106. doi: 10.1139/gen-2012-0106 URL
[7]	Ishikawa S, Abe T, Kuramata M, Yamaguchi M, Ando T, Yamamoto T, Yano M. A major quantitative trait locus for increasing cadmium-specific concentration in rice grain is located on the short arm of chromosome 7[J]. Journal of Experimental Botany, 2009, 61(3):923-934.doi: 10.1093/jxb/erp360. doi: 10.1093/jxb/erp360 URL
[8]	骆旭添. 水稻苗期耐镉胁迫的QTL定位及其与环境互作效应分析[D]. 福州: 福建农林大学, 2007.doi: 10.7666/d.y2045758. doi: 10.7666/d.y2045758
	Luo X T. QTL mapping for seedling Cd tolerance in rice (Oryza sativa L.)and analysis of QTL × environment interaction[D]. Fuzhou: Fujian Agriculture and Forestry University, 2007.
[9]	Pan X W, Li Y C, Liu W Q, Liu S X, Min J, Xiong H B, Dong Z, Duan Y H, Yu Y Y, Li X X. QTL mapping and candidate gene analysis of cadmium accumulation in polished rice by genome-wide association study[J]. Scientific Reports, 2020, 10:11791.doi: 10.1038/s41598-020-68742-4. doi: 10.1038/s41598-020-68742-4 pmid: 32678216
[10]	Li C S, Wang P, Wu G L, Wang Y N, Cheng Q, Cai Y C, Zhou D H, Li C J, Zhang X Y, Tan J G, Li C J, He H H, Bian J M. Additive and epistatic QTL on cadmium(Cd)tolerance associated with seed germinating ability in rice[J]. Journal of Plant Growth Regulation, 2021, 40(5):2115-2123.doi: 10.1007/s00344-020-10258-2. doi: 10.1007/s00344-020-10258-2 URL
[11]	Ueno D, Kono I, Yokosho K, Ando T, Yano M, Ma J F. A major quantitative trait locus controlling cadmium translocation in rice(Oryza sativa)[J]. The New Phytologist, 2009, 182(3):644-653.doi: 10.1111/j.1469-8137.2009.02784.x. doi: 10.1111/j.1469-8137.2009.02784.x URL
[12]	Luo J S, Huang J, Zeng D L, Peng J S, Zhang G B, Ma H L, Guan Y, Yi H Y, Fu Y L, Han B, Lin H X, Qian Q, Gong J M. A defensin-like protein drives cadmium efflux and allocation in rice[J]. Nature Communications, 2018, 9(1):645.doi: 10.1038/s41467-018-03088-0. doi: 10.1038/s41467-018-03088-0 URL
[13]	Tian S Q, Liang S, Qiao K, Wang F H, Zhang Y X, Chai T Y. Co-expression of multiple heavy metal transporters changes the translocation,accumulation,and potential oxidative stress of Cd and Zn in rice(Oryza sativa)[J]. Journal of Hazardous Materials, 2019, 380:120853.doi: 10.1016/j.jhazmat.2019.120853. doi: 10.1016/j.jhazmat.2019.120853 URL
[14]	Chen J, Huang X Y, Salt D E, Zhao F J. Mutation in OsCADT1 enhances cadmium tolerance and enriches selenium in rice grain[J]. The New Phytologist, 2020, 226(3):838-850.doi: 10.1111/nph.16404. doi: 10.1111/nph.16404 URL
[15]	Shimo H, Ishimaru Y, An G, Yamakawa T, Nakanishi H, Nishizawa N K. Low cadmium(LCD),a novel gene related to cadmium tolerance and accumulation in rice[J]. Journal of Experimental Botany, 2011, 62(15):5727-5734.doi: 10.1093/jxb/err300. doi: 10.1093/jxb/err300 pmid: 21908474
[16]	Yan L, Chen C, Zhu Y C, La Y P, Zhang S J, Ding G Y, Qu J J. Cadmium-induced phytotoxicity and tolerance response in the low-Cd accumulator of Chinese cabbage(Brassica pekinensis L.)seedlings[J]. International Journal of Phytoremediation, 2021, 23(13):1365-1375.doi: 10.1080/15226514.2021.1897778. doi: 10.1080/15226514.2021.1897778 URL
[17]	李炜星, 欧阳林娟, 文文, 熊玉毅, 徐伟清, 彭小松, 陈小荣, 贺晓鹏, 傅军如, 边建民, 徐杰, 周大虎, 贺浩华, 孙晓棠, 朱昌兰. 水稻幼苗耐镉胁迫QTL的定位研究[J]. 江西农业大学学报, 2019, 41(1):19-24.doi: 10.13836/j.jjau.2019003. doi: 10.13836/j.jjau.2019003
	Li W X, Ouyang L J, Wen W, Xiong Y Y, Xu W Q, Peng X S, Chen X R, He X P, Fu J R, Bian J M, Xu J, Zhou D H, He H H, Sun X T, Zhu C L. Identification of QTL for cadmium tolerance at seedling stage of rice(Oryza sativa L.)[J]. Acta Agriculturae Universitatis Jiangxiensis, 2019, 41(1):19-24.
[18]	黄诗颖, 谭景艾, 王鹏, 蒋宁飞, 贺浩华, 边建民. 以回交重组自交系定位水稻萌芽期耐镉胁迫相关QTLs[J]. 生物技术进展, 2021, 11(2):176-181.doi: 10.19586/j.2095-2341.2020.0135. doi: 10.19586/j.2095-2341.2020.0135
	Huang S Y, Tan J G, Wang P, Jiang N F, He H H, Bian J M. Mapping QTLs related to tolerance to cadmium in rice during germination by backcross recombinant inbred lines[J]. Current Biotechnology, 2021, 11(2):176-181.
[19]	赵均良, 张少红, 杨梯丰, 董景芳, 刘清, 付华, 毛兴学, 刘斌. 181份多样性籼稻种质苗期和成熟期镉积累表型评价[J]. 分子植物育种, 2018, 16(18):6080-6087.doi: 10.13271/j.mpb.016.006080. doi: 10.13271/j.mpb.016.006080
	Zhao J L, Zhang S H, Yang T F, Dong J F, Liu Q, Fu H, Mao X X, Liu B. Phenotype evaluation of Cd accumulation of 181 diverse indica germplasm at seedling and mature stages[J]. Molecular Plant Breeding, 2018, 16(18):6080-6087.
[20]	Liu X Y, Chen S L, Chen M X, Zheng G Y, Peng Y, Shi X L, Qin P, Xu X Y, Teng S. Association study reveals genetic loci responsible for arsenic,cadmium and lead accumulation in rice grain in contaminated farmlands[J]. Frontiers in Plant Science, 2019, 10:61.doi: 10.3389/fpls.2019.00061. doi: 10.3389/fpls.2019.00061 URL
[21]	Zhao J, He Y Q, Li X Y, Weng X N, Feng D F, Ying J F, Wang Z F. An integrated RNA-Seq and physiological study reveals gene responses involving in the initial imbibition of seed germination in rice[J]. Plant Growth Regulation, 2020, 90(2):249-263.doi: 10.1007/s10725-019-00567-2. doi: 10.1007/s10725-019-00567-2 URL
[22]	Lei L, Zheng H L, Bi Y L, Yang L M, Liu H L, Wang J G, Sun J, Zhao H W, Li X W, Li J M, Lai Y C, Zou D T. Identification of a major QTL and candidate gene analysis of salt tolerance at the bud burst stage in rice(Oryza sativa L.)using QTL-seq and RNA-seq[J]. Rice, 2020, 13(1):55.doi: 10.1186/s12284-020-00416-1. doi: 10.1186/s12284-020-00416-1 pmid: 32778977
[23]	Wei H B, Lou Q J, Xu K, Zhou L G, Chen S J, Chen L, Luo L J. Pattern of alternative splicing different associated with difference in rooting depth in rice[J]. Plant and Soil, 2020, 449(1/2):233-248.doi: 10.1007/s11104-020-04451-1. doi: 10.1007/s11104-020-04451-1 URL
[24]	Ereful N C, Liu L Y, Greenland A, Powell W, MacKay I, Leung H. RNA-seq reveals differentially expressed genes between two indica inbred rice genotypes associated with drought-yield QTLs[J]. Agronomy, 2020, 10(5):621.doi: 10.3390/agronomy10050621. doi: 10.3390/agronomy10050621 URL
[25]	Chen L, Shi S L, Jiang N F, Khanzada H, Wassan G M, Zhu C L, Peng X S, Xu J, Chen Y J, Yu Q Y, He X P, Fu J R, Chen X R, Hu L F, Ouyang L J, Sun X T, He H H, Bian J M. Genome-wide analysis of long non-coding RNAs affecting roots development at an early stage in the rice response to cadmium stress[J]. BMC Genomics, 2018, 19(1):460.doi: 10.1186/s12864-018-4807-6. doi: 10.1186/s12864-018-4807-6 pmid: 29902991
[26]	Yang J, Guo Z H, Luo L X, Gao Q L, Xiao W M, Wang J F, Wang H, Chen Z Q, Guo T. Identification of QTL and candidate genes involved in early seedling growth in rice via high-density genetic mapping and RNA-seq[J]. The Crop Journal, 2021, 9(2):360-371.doi: 10.1016/j.cj.2020.08.010. doi: 10.1016/j.cj.2020.08.010 URL
[27]	Mortier V, Fenta B A, Martens C, Rombauts S, Holsters M, Kunert K, Goormachtig S. Search for nodulation-related CLE genes in the genome of Glycine max[J]. Journal of Experimental Botany, 2011, 62(8):2571-2583.doi: 10.1093/jxb/erq426. doi: 10.1093/jxb/erq426 URL
[28]	Chen Z, Gallie D R. Increasing tolerance to ozone by elevating foliar ascorbic acid confers greater protection against ozone than increasing avoidance[J]. Plant Physiology, 2005, 138(3):1673-1689.doi: 10.1104/pp.105.062000. doi: 10.1104/pp.105.062000 pmid: 15951482
[29]	Neill S O, Gould K S, Kilmartin P A, Mitchell K A, Markham K R. Antioxidant activities of red versus green leaves in Elatostema rugosum[J]. Plant Cell and Environment, 2002, 25:539-547.doi: 10.1046/J.1365-3040.2002.00837.X. doi: 10.1046/J.1365-3040.2002.00837.X URL
[30]	Roychoudhury A, Basu S, Sengupta D N. Antioxidants and stress-related metabolites in the seedlings of two indica rice varieties exposed to cadmium chloride toxicity[J]. Acta Physiologiae Plantarum, 2012, 34(3):835-847.doi: 10.1007/s11738-011-0881-y. doi: 10.1007/s11738-011-0881-y URL
[31]	Bai B, Wu J, Sheng W T, Zhou B, Zhou L J, Zhuang W, Yao D P, Deng Q Y. Comparative analysis of anther transcriptome profiles of two different rice male sterile lines genotypes under cold stress[J]. International Journal of Molecular Sciences, 2015, 16(5):11398-11416.doi: 10.3390/ijms160511398. doi: 10.3390/ijms160511398 pmid: 25993302
[32]	Shin J H, Yoshimoto K, Ohsumi Y, Jeon J S, An G. OsATG10b,an autophagosome component,is needed for cell survival against oxidative stresses in rice[J]. Molecules and Cells, 2009, 27(1):67-74.doi: 10.1007/s10059-009-0006-2. doi: 10.1007/s10059-009-0006-2 URL
[33]	Xia K F, Liu T, Ouyang J, Wang R, Fan T, Zhang M Y. Genome-wide identification,classification,and expression analysis of autophagy-associated gene homologues in rice(Oryza sativa L.)[J]. DNA Research, 2011, 18(5):363-377.doi: 10.1093/dnares/dsr024. doi: 10.1093/dnares/dsr024 URL
[34]	Ammar W B, Nouairi I, Zarrouk M, Jemal F. The effect of cadmium on lipid and fatty acid biosynthesis in tomato leaves[J]. Biologia, 2008, 63(1):86-93.doi: 10.2478/s11756-008-0002-6. doi: 10.2478/s11756-008-0002-6 URL
[35]	Lin C Y, Trinh N N, Fu S F, Hsiung Y C, Chia L C, Lin C W, Huang H J. Comparison of early transcriptome responses to copper and cadmium in rice roots[J]. Plant Molecular Biology, 2013, 81(4/5):507-522.doi: 10.1007/s11103-013-0020-9. doi: 10.1007/s11103-013-0020-9 URL

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