木薯重要性状基因的研究进展

doi:10.13560/j.cnki.biotech.bull.1985.2022-1274

摘要/Abstract

摘要：

木薯是全球热区的重要粮食作物、经济作物和能源作物，但其生物学研究和育种进展落后于主要粮食作物，分子育种是木薯遗传改良的重要驱动力，挖掘木薯重要性状相关基因是实现其传统育种向分子育种转变的基础和前提。本文系统地总结了木薯株型、产量、品质、抗逆等性状基因以及相关基因功能表征的最新进展，并指出构建自交分离群体和多组学整合是未来挖掘木薯关键基因的重要手段。本文拟为推进功能基因组研究成果应用于木薯育种技术体系建设提供参考，为木薯遗传改良提供理论指导。

关键词: 木薯, 重要性状, 基因挖掘, 分子标记, 功能基因

Abstract:

Cassava（Manihot esculenta Crantz）is an important food crop, cash crop and energy crop in global tropical regions, but the biology research and breeding progress have lagged behind major food crops. Molecular breeding is the important driving force for cassava genetic improvement. Discovery the genes related to important traits is the foundation and premise for the transformation from traditional breeding to molecular breeding. In this paper, systematically, we recapitulate recent progress on the genes related to the traits, such as plant morphology, yield, tuber quality, stress resistance, in cassava, also functional characterization of some genes. We further point out that both of constructing self-crossing population and multiple-omics data integrated are important ways to discovery the key genes in cassava in the future. This paper aims to provide reference for promoting the application of results of functional genome studies in the construction of cassava breeding technology system, and provide theoretical guidance for cassava genetic improvement.

Key words: cassava, important trait, gene discovery, molecular marker, functional gene

肖亮, 吴正丹, 陆柳英, 施平丽, 尚小红, 曹升, 曾文丹, 严华兵. 木薯重要性状基因的研究进展[J]. 生物技术通报, 2023, 39(6): 31-48.

XIAO Liang, WU Zheng-dan, LU Liu-ying, SHI Ping-li, SHANG Xiao-hong, CAO Sheng, ZENG Wen-dan, YAN Hua-bing. Research Progress of Important Traits Genes in Cassava[J]. Biotechnology Bulletin, 2023, 39(6): 31-48.

图/表 2

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性状 Trait	基因ID Gene ID	基因注释 Gene annotation	参考文献Reference
第一分支高度 The first branch height	Manes.09G041800	钙调素蛋白激酶20 Calcium-dependent protein kinase 20	[12]
	Manes.09G041900	ABC-2型转运蛋白 ABC-2 type transporter protein
	Manes.02G212100	果胶酸酯裂解酶超家族蛋白 Pectin lyase-like superfamily protein
	Manes.02G212200	未知蛋白 Unknown protein
	Manes.03G061100	半胱氨酸氨基转移 Cysteine aminotransferase
	Manes.03G061200	四跨膜蛋白 Tetraspanin
	Manes.10G123100	跨膜蛋白9超家族成员1 Transmembrane 9 superfamily member 1
	Manes.09G143900	未知蛋白 Unknown protein
	Manes.09G144000	未知蛋白 Unknown protein
	Manes.09G143800	NAC转录因子87 NAC transcriptional factor 87
	Manes.03G059800	未知蛋白 Unknown protein
茎直径 Stem diameter	Manes.03G059800	未知蛋白 Unknown protein	[12]
	Manes.03G170000	加工rRNA蛋白EFG1 rRNA-processing protein EFG1
	Manes.03G169900	α-甘露糖苷酶 alpha-mannosidase
	Manes.14G021000	NADH脱氢酶-1-α-亚基复合体亚基5 NADH dehydrogenase-1-alpha-subcomplex subunit 5
	Manes.14G020900	SNARE-like蛋白 SNARE-like protein
	Manes.07G117800	POTUNDIFOLIA蛋白 POTUNDIFOLIA protein
	Manes.07G117700	未知蛋白 Unknown protein
	Manes.07G007700	细胞色素b5域RLF Cytochrome b5 domain-containing protein RLF
	Manes.07G007800	环阿屯醇-碳-24-甲基转移酶 Cycloartenol-C-24-methyltransferase
	Manes.06G047300	未知蛋白 Unknown protein
叶指宽 Lobular width	Manes.05G164600	核糖体蛋白S9 Ribosomal protein S9	[12]
	Manes.05G164500	多药物及有毒化合物外排转运蛋白 Multidrug and toxic compound extrusion
	Manes.03G156800	线粒体编辑因子22 Mitochondrial editing factor 22
	Manes.03G156900	MYB转录因子 MYB transcriptional factor
	Manes.03G157000	膜蛋白PM19L Membrane protein PM19L
	Manes.10G312000	OBF3蛋白 OBF3 protein
叶指长 Lobular length	Manes.05G026500	葡萄糖-6-磷酸差相异构酶 Glucose-6-phosphate1-epimerase	[12]
	Manes.05G026600	未知蛋白 Unknown protein
	Manes.05G026700	G-box结合因子 G-box binding factor
	Manes.06G034000	26S蛋白酶体调节亚基 26S proteasome regulatory subunit
	Manes.18G018500	组氨酸超家族蛋白 Histone superfamily protein
	Manes.04G057300	醛氧化酶2 Aldehyde oxidase 2
叶柄长度 Petiole length	Manes.01G201900	bHLH转录因子 bHLH transcriptional factor	[12]
叶柄长度 Petiole length	Manes.01G202000	未知蛋白 Unknown protein	[12]
叶指长宽比 Leaf aspect ratio	Manes.03G156800	线粒体编辑因子22 Mitochondrial editing factor 22	[12]
	Manes.03G156900	MYB转录因子 MYB transcriptional factor
	Manes.03G157000	AWPM-19-like家族蛋白 AWPM-19-like family protein
	Manes.06G034000	26S蛋白酶体调节亚基 26S proteasome regulatory subunit
	Manes.14G056200	MA3蛋白 MA3 protein
	Manes.14G056100	MA3蛋白 MA3 protein
	Manes.14G056300	未知蛋白 Unknown protein
	Manes.01G182700	蔗糖转化酶/果胶甲基转移酶抑制子 Invertase/pectin methylesterase inhibitor
	Manes.01G182800	未知蛋白 Unknown protein
	Manes.01G182900	甘氨酸富集蛋白 Glycine-rich protein
	Manes.03G184600	P-loop-核苷三磷酸水解酶 P-loop-containing nucleoside triphosphate hydrolase
	Manes.03G186500	α-水解酶 α-hydrolase
	Manes.10G031200	OBF3蛋白 OBF3 protein
叶形 The shape of leaf	Manes.15G136200	KNOX1和KNOX2蛋白 KNOX1 and KNOX2 protein	[12]
叶柄颜色 Petiole color	Manes.01G115400	MYB转录因子6 MYB transcriptional factor 6	[13]
绿叶 Green leaf	Manes.01G115400	MYB转录因子6 MYB transcriptional factor 6	[13]
干物质重量 Dry mass weight	Manes.02G192500	未知蛋白 Unknown protein	[12]
	Manes.02G192600	RING/U-box锌指家族蛋白 RING/U-box zinc finger family protein
	Manes.02G169700	结瘤MtN3家族蛋白 Nodulin MtN3 family protein
	Manes.02G169800	未知蛋白 Unknown protein
	Manes.02G154700	β-D-木糖苷酶7 Beta-D-xylosidase 7
	Manes.02G154800	类钙神经素金属磷酸酯酶 Calcineurin-like metallo-phosphoesterase
储藏根数目 Number of storage roots	Manes.05G125100	丝氨酸/苏氨酸蛋白激酶IRE1b Ser/Thr protein kinase IRE1b	[12]
	Manes.05G125200	3-磷酸甘油醛脱氢酶B亚基 Glyceraldehyde-3-phosphate dehydrogenase B subunit
	Manes.05G125300	未知蛋白 Unknown protein
	Manes.04G057900	未知蛋白 Unknown protein
	Manes.04G058000	未知蛋白 Unknown protein
	Manes.09G099100	未知蛋白 Unknown protein
	Manes.09G099200	铜转运蛋白6 Copper transport protein 6
	Manes.09G099300	热激蛋白HSP20-like Heat shock protein HSP20-like
储藏根重量 Storage roots weight	Manes.02G169700	结瘤MtN3家族蛋白 Nodulin MtN3 family protein	[12]
储藏根重量 Storage roots weight	Manes.02G169800	未知蛋白 Unknown protein	[12]
收获指数 Harvest index	Manes.02G035900	β-果糖基转移酶 Beta-fructofuranosidase	[13]
收获指数 Harvest index	Manes.02G037700	4 glucan phosphorylase L isozyme	[13]
淀粉含量 Starch content	Manes.13G023300	β-1,4-氮-乙酰葡糖氨基转移酶 Beta-1,4-N-acetylglucosaminyl transferase	[12]
	Manes.13G023400	NAD（P）-氧化还原酶 NAD（P）-linked oxidoreductase
	Manes.05G177800	碳水化合物酯酶 Carbohydrate esterase
总类胡萝卜素含量 Total carotenoid content	Manes.01G124200	八氢番茄红素酶2 Phytoene synthase 2	[13⇓⇓-16]
	Manes.05G051700	β-胡萝卜素双加氧酶 Beta-carotene dioxgenese	[13]
	Manes.15G102000	β-胡萝卜素双加氧酶 Beta-carotene dioxgenese	[13]
	Manes.16G099600	β-番茄红素环化酶 Lycopene beta cyclase	[16]
	Manes.09G008200	ε-番茄红素E环化酶 Lycopene epsilon cyclase
	Manes.06G152200	β-胡萝卜素羟化酶 Beta-carotene hydroxylase
干物质含量 Dry mass content	Manes.14G007500	质体3-磷酸甘油醛脱氢酶2 Glyceraldehyde-3-phosphate dehydrogenase of plastid 2	[12]
	Manes.16G000700	ATP酶WRNIP1 ATPase WRNIP1
	Manes.16G000800	亮氨酸富集蛋白受体激酶 Leucine rich repeat receptor like kinase
	Manes.01G123000	UDP-葡萄糖焦磷酸化酶 UTP-glucose pyrophosphorylase	[13,17]
	Manes.01G123800	蔗糖合酶 Sucrose synthase	[13,17]
	Manes.06G103600	双向糖转运蛋白SWEET5 Bidirectional sugar transporter SWEET5	[13]
	Manes.15G011300	RAG1激活蛋白 RAG1 activating protein
	Manes.16G109200	解螺旋酶 Helicase
氢氰酸苷含量 HCN content	Manes.16G007900	多药物及有毒化合物外排转运蛋白 Multidrug and toxic compound extrusion	[18]
CMD2抗性 CMD2 resistance	Manes.12G076200	过氧化物酶 Peroxidase	[13]
	Manes.12G076300	过氧化物酶 Peroxidase
	Manes.14G058400	TCP家族转录因子 TCP family transcriptional factor
CBSD抗性 CBSD resistance	Manes.11G130500	甘氨酸富集蛋白 Glycine-rich repeat protein	[19]
	Manes.11G130000	亮氨酸富集蛋白Leucine-rich repeat protein
	Manes.11G130200	触发因子分子伴侣 Trigger factor chaperone
	Manes.11G131100	U-box蛋白33 U-box domain-containing protein 33
	Manes.11G127100	甘氨酸富集蛋白 Glycine-rich repeat protein	[20]
	Cassava4.1_019379m	LysM结构域包含蛋白 LysM domain containing protein
	Cassava4.1_00037m	3.5.2.9-5-羟脯氨酸酶 3.5.2.9-5-oxoprolinase enzyme
CGM抗性CGM resistance	Manes.08G058000	MYB转录因子106 MYB transcriptional factor 106	[13,21]
	Manes.08G045400	MYB-like螺旋-转角-螺旋转录因子 MYB-like helix-turn-helix transcriptional factor	[21]
	Manes.08G058500	C2H2-like锌指转录因子 C2H2-like Zn finger transcriptional factor
	Manes.08G048200	C2H2-type锌指转录因子 C2H2-type Zn finger transcriptional factor
	Manes.08G048800	富含ARM重复域CCCH锌指转录因子 CCCH-type Zn finger transcriptional factor with ARM repeat domain
	Manes.08G034200	Dof-type锌指转录因子 Dof-type Zn finger transcriptional factor
	Manes.08G046400	K同源域CCCH-type锌指转录因子 K homology-domain-containing protein-/-Zn finger（CCCH-type）transcriptional factor
	Manes.08G041900	锌指转录因子8 Zn finger transcriptional factor 8
	Manes.08G035100	MADS-box转录因子AGAMOUS-like 80 MADS-box transcriptional factor AGAMOUS-like 80
	Manes.08G043900	同源框亮氨酸拉链蛋白HOX11 Homeobox-leucine zipper protein HOX11
	Manes.08G024700	亮氨酸拉链转录因子 Basic leucine zipper transcriptional factor
	Manes.08G046700	亮氨酸拉链转录因子 Basic leucine zipper transcriptional factor
	Manes.08G026900	SAUR-like生长素响应因子 SAUR-like auxin-responsive factor
	Manes.08G026500	三角状五肽重复蛋白 Pentatricopeptide repeat protein
	Manes.08G053900	三角状五肽重复蛋白 Pentatricopeptide repeat protein
	Manes.08G060500	三角状五肽重复蛋白 Pentatricopeptide repeat protein
	Manes.08G044000	表皮毛双折射相关蛋白 Trichome birefringence-like protein

性状 Trait	基因ID Gene ID	基因注释 Gene annotation	参考文献Reference
第一分支高度 The first branch height	Manes.09G041800	钙调素蛋白激酶20 Calcium-dependent protein kinase 20	[12]
	Manes.09G041900	ABC-2型转运蛋白 ABC-2 type transporter protein
	Manes.02G212100	果胶酸酯裂解酶超家族蛋白 Pectin lyase-like superfamily protein
	Manes.02G212200	未知蛋白 Unknown protein
	Manes.03G061100	半胱氨酸氨基转移 Cysteine aminotransferase
	Manes.03G061200	四跨膜蛋白 Tetraspanin
	Manes.10G123100	跨膜蛋白9超家族成员1 Transmembrane 9 superfamily member 1
	Manes.09G143900	未知蛋白 Unknown protein
	Manes.09G144000	未知蛋白 Unknown protein
	Manes.09G143800	NAC转录因子87 NAC transcriptional factor 87
	Manes.03G059800	未知蛋白 Unknown protein
茎直径 Stem diameter	Manes.03G059800	未知蛋白 Unknown protein	[12]
	Manes.03G170000	加工rRNA蛋白EFG1 rRNA-processing protein EFG1
	Manes.03G169900	α-甘露糖苷酶 alpha-mannosidase
	Manes.14G021000	NADH脱氢酶-1-α-亚基复合体亚基5 NADH dehydrogenase-1-alpha-subcomplex subunit 5
	Manes.14G020900	SNARE-like蛋白 SNARE-like protein
	Manes.07G117800	POTUNDIFOLIA蛋白 POTUNDIFOLIA protein
	Manes.07G117700	未知蛋白 Unknown protein
	Manes.07G007700	细胞色素b5域RLF Cytochrome b5 domain-containing protein RLF
	Manes.07G007800	环阿屯醇-碳-24-甲基转移酶 Cycloartenol-C-24-methyltransferase
	Manes.06G047300	未知蛋白 Unknown protein
叶指宽 Lobular width	Manes.05G164600	核糖体蛋白S9 Ribosomal protein S9	[12]
	Manes.05G164500	多药物及有毒化合物外排转运蛋白 Multidrug and toxic compound extrusion
	Manes.03G156800	线粒体编辑因子22 Mitochondrial editing factor 22
	Manes.03G156900	MYB转录因子 MYB transcriptional factor
	Manes.03G157000	膜蛋白PM19L Membrane protein PM19L
	Manes.10G312000	OBF3蛋白 OBF3 protein
叶指长 Lobular length	Manes.05G026500	葡萄糖-6-磷酸差相异构酶 Glucose-6-phosphate1-epimerase	[12]
	Manes.05G026600	未知蛋白 Unknown protein
	Manes.05G026700	G-box结合因子 G-box binding factor
	Manes.06G034000	26S蛋白酶体调节亚基 26S proteasome regulatory subunit
	Manes.18G018500	组氨酸超家族蛋白 Histone superfamily protein
	Manes.04G057300	醛氧化酶2 Aldehyde oxidase 2
叶柄长度 Petiole length	Manes.01G201900	bHLH转录因子 bHLH transcriptional factor	[12]
叶柄长度 Petiole length	Manes.01G202000	未知蛋白 Unknown protein	[12]
叶指长宽比 Leaf aspect ratio	Manes.03G156800	线粒体编辑因子22 Mitochondrial editing factor 22	[12]
	Manes.03G156900	MYB转录因子 MYB transcriptional factor
	Manes.03G157000	AWPM-19-like家族蛋白 AWPM-19-like family protein
	Manes.06G034000	26S蛋白酶体调节亚基 26S proteasome regulatory subunit
	Manes.14G056200	MA3蛋白 MA3 protein
	Manes.14G056100	MA3蛋白 MA3 protein
	Manes.14G056300	未知蛋白 Unknown protein
	Manes.01G182700	蔗糖转化酶/果胶甲基转移酶抑制子 Invertase/pectin methylesterase inhibitor
	Manes.01G182800	未知蛋白 Unknown protein
	Manes.01G182900	甘氨酸富集蛋白 Glycine-rich protein
	Manes.03G184600	P-loop-核苷三磷酸水解酶 P-loop-containing nucleoside triphosphate hydrolase
	Manes.03G186500	α-水解酶 α-hydrolase
	Manes.10G031200	OBF3蛋白 OBF3 protein
叶形 The shape of leaf	Manes.15G136200	KNOX1和KNOX2蛋白 KNOX1 and KNOX2 protein	[12]
叶柄颜色 Petiole color	Manes.01G115400	MYB转录因子6 MYB transcriptional factor 6	[13]
绿叶 Green leaf	Manes.01G115400	MYB转录因子6 MYB transcriptional factor 6	[13]
干物质重量 Dry mass weight	Manes.02G192500	未知蛋白 Unknown protein	[12]
	Manes.02G192600	RING/U-box锌指家族蛋白 RING/U-box zinc finger family protein
	Manes.02G169700	结瘤MtN3家族蛋白 Nodulin MtN3 family protein
	Manes.02G169800	未知蛋白 Unknown protein
	Manes.02G154700	β-D-木糖苷酶7 Beta-D-xylosidase 7
	Manes.02G154800	类钙神经素金属磷酸酯酶 Calcineurin-like metallo-phosphoesterase
储藏根数目 Number of storage roots	Manes.05G125100	丝氨酸/苏氨酸蛋白激酶IRE1b Ser/Thr protein kinase IRE1b	[12]
	Manes.05G125200	3-磷酸甘油醛脱氢酶B亚基 Glyceraldehyde-3-phosphate dehydrogenase B subunit
	Manes.05G125300	未知蛋白 Unknown protein
	Manes.04G057900	未知蛋白 Unknown protein
	Manes.04G058000	未知蛋白 Unknown protein
	Manes.09G099100	未知蛋白 Unknown protein
	Manes.09G099200	铜转运蛋白6 Copper transport protein 6
	Manes.09G099300	热激蛋白HSP20-like Heat shock protein HSP20-like
储藏根重量 Storage roots weight	Manes.02G169700	结瘤MtN3家族蛋白 Nodulin MtN3 family protein	[12]
储藏根重量 Storage roots weight	Manes.02G169800	未知蛋白 Unknown protein	[12]
收获指数 Harvest index	Manes.02G035900	β-果糖基转移酶 Beta-fructofuranosidase	[13]
收获指数 Harvest index	Manes.02G037700	4 glucan phosphorylase L isozyme	[13]
淀粉含量 Starch content	Manes.13G023300	β-1,4-氮-乙酰葡糖氨基转移酶 Beta-1,4-N-acetylglucosaminyl transferase	[12]
	Manes.13G023400	NAD（P）-氧化还原酶 NAD（P）-linked oxidoreductase
	Manes.05G177800	碳水化合物酯酶 Carbohydrate esterase
总类胡萝卜素含量 Total carotenoid content	Manes.01G124200	八氢番茄红素酶2 Phytoene synthase 2	[13⇓⇓-16]
	Manes.05G051700	β-胡萝卜素双加氧酶 Beta-carotene dioxgenese	[13]
	Manes.15G102000	β-胡萝卜素双加氧酶 Beta-carotene dioxgenese	[13]
	Manes.16G099600	β-番茄红素环化酶 Lycopene beta cyclase	[16]
	Manes.09G008200	ε-番茄红素E环化酶 Lycopene epsilon cyclase
	Manes.06G152200	β-胡萝卜素羟化酶 Beta-carotene hydroxylase
干物质含量 Dry mass content	Manes.14G007500	质体3-磷酸甘油醛脱氢酶2 Glyceraldehyde-3-phosphate dehydrogenase of plastid 2	[12]
	Manes.16G000700	ATP酶WRNIP1 ATPase WRNIP1
	Manes.16G000800	亮氨酸富集蛋白受体激酶 Leucine rich repeat receptor like kinase
	Manes.01G123000	UDP-葡萄糖焦磷酸化酶 UTP-glucose pyrophosphorylase	[13,17]
	Manes.01G123800	蔗糖合酶 Sucrose synthase	[13,17]
	Manes.06G103600	双向糖转运蛋白SWEET5 Bidirectional sugar transporter SWEET5	[13]
	Manes.15G011300	RAG1激活蛋白 RAG1 activating protein
	Manes.16G109200	解螺旋酶 Helicase
氢氰酸苷含量 HCN content	Manes.16G007900	多药物及有毒化合物外排转运蛋白 Multidrug and toxic compound extrusion	[18]
CMD2抗性 CMD2 resistance	Manes.12G076200	过氧化物酶 Peroxidase	[13]
	Manes.12G076300	过氧化物酶 Peroxidase
	Manes.14G058400	TCP家族转录因子 TCP family transcriptional factor
CBSD抗性 CBSD resistance	Manes.11G130500	甘氨酸富集蛋白 Glycine-rich repeat protein	[19]
	Manes.11G130000	亮氨酸富集蛋白Leucine-rich repeat protein
	Manes.11G130200	触发因子分子伴侣 Trigger factor chaperone
	Manes.11G131100	U-box蛋白33 U-box domain-containing protein 33
	Manes.11G127100	甘氨酸富集蛋白 Glycine-rich repeat protein	[20]
	Cassava4.1_019379m	LysM结构域包含蛋白 LysM domain containing protein
	Cassava4.1_00037m	3.5.2.9-5-羟脯氨酸酶 3.5.2.9-5-oxoprolinase enzyme
CGM抗性CGM resistance	Manes.08G058000	MYB转录因子106 MYB transcriptional factor 106	[13,21]
	Manes.08G045400	MYB-like螺旋-转角-螺旋转录因子 MYB-like helix-turn-helix transcriptional factor	[21]
	Manes.08G058500	C2H2-like锌指转录因子 C2H2-like Zn finger transcriptional factor
	Manes.08G048200	C2H2-type锌指转录因子 C2H2-type Zn finger transcriptional factor
	Manes.08G048800	富含ARM重复域CCCH锌指转录因子 CCCH-type Zn finger transcriptional factor with ARM repeat domain
	Manes.08G034200	Dof-type锌指转录因子 Dof-type Zn finger transcriptional factor
	Manes.08G046400	K同源域CCCH-type锌指转录因子 K homology-domain-containing protein-/-Zn finger（CCCH-type）transcriptional factor
	Manes.08G041900	锌指转录因子8 Zn finger transcriptional factor 8
	Manes.08G035100	MADS-box转录因子AGAMOUS-like 80 MADS-box transcriptional factor AGAMOUS-like 80
	Manes.08G043900	同源框亮氨酸拉链蛋白HOX11 Homeobox-leucine zipper protein HOX11
	Manes.08G024700	亮氨酸拉链转录因子 Basic leucine zipper transcriptional factor
	Manes.08G046700	亮氨酸拉链转录因子 Basic leucine zipper transcriptional factor
	Manes.08G026900	SAUR-like生长素响应因子 SAUR-like auxin-responsive factor
	Manes.08G026500	三角状五肽重复蛋白 Pentatricopeptide repeat protein
	Manes.08G053900	三角状五肽重复蛋白 Pentatricopeptide repeat protein
	Manes.08G060500	三角状五肽重复蛋白 Pentatricopeptide repeat protein
	Manes.08G044000	表皮毛双折射相关蛋白 Trichome birefringence-like protein

基因名称 Gene name	功能 Function	产物 Product	生理机制 Physiological mechanism	参考文献 Reference
MeCWINV3	负调控淀粉积累 Negative regulation of starch accumulation	细胞壁转化酶 Cell wall invertases	抑制叶片中蔗糖向块根转移Inhibited sugar export from leaves to storage roots	[25]
MeSBE1	负调控直链淀粉形成 Negative regulation of high-amylose production	淀粉分支酶1 Starch branch enzyme 1	促进块根中支链淀粉链长分布 Promoted chain-length distribution of amylopectin	[26]
MeSBE2	负调控直链淀粉形成 Negative regulation of high-amylose production	淀粉分支酶2 Starch branch enzyme 2	促进块根中支链淀粉链长分布 Promoted chain-length distribution of amylopectin	[26]
MeGBSSI	正调控糯木薯形成 Positive regulation of waxy cassava	颗粒结合淀粉合酶I Granule-bound starch synthase I	合成蜡质淀粉 Synthesized waxy starch	[27]
MePSY2	正调控黄色薯肉形成 Positive regulation of yellow tuber	八氢番茄红素酶2 Phytoene synthase 2	促进β-胡萝卜素合成 Promoted β-carotene accumulation	[14]
MeCAT1 （Co-expressed with MeCu/ZnSOD）	延缓PPD Delay PPD	过氧化氢酶 Catalase	强化ROS清除 Enhanced ROS scavenging	[28]
HNL	负调控块根中HCN含量 Negative regulation of HCN content in tuber	羟基腈裂解酶Hydroxynitrile lyase	促进丙酮氰醇分解 Promoted the decomposition of acetone cyanol	[29]
MeDREB1D	正调控抗旱 Positive regulation of drought resistance	AP2转录因子 AP2 transcriptional factor	降低MDA含量，并可能强化了ROS清除 Decreased MDA content, and may enhance ROS scavenging	[30]
MeDREB1D	正调控抗寒 Positive regulation of cold resistance	AP2转录因子 AP2 transcriptional factor	可能强化了ROS清除 May enhance ROS scavenging	[30]
MeDREB1A	正调控抗旱 Positive regulation of drought resistance	AP2转录因子 AP2 transcriptional factor	增加脯氨酸含量 Increased proline content	[31]
MeDREB1A	正调控抗寒 Positive regulation of cold resistance	AP2转录因子 AP2 transcriptional factor	增加脯氨酸含量 Increased proline content	[31]
MeRAV5	正调控抗旱 Positive regulation of drought resistance	AP2转录因子 AP2 transcriptional factor	降低H₂O₂含量和促进木质素积累 Decreased H₂O₂ content and promote lignin accumulation	[32]
MeGRX360	正调控抗旱 Positive regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	降低H₂O₂含量 Decreased H₂O₂content	[33]
MeGRX058	正调控抗旱 Positive regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	降低H₂O₂含量 Decreased H₂O₂content
MeGRX785	负调控抗旱 Negative regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	降低H₂O₂含量 Decreased H₂O₂content
MeGRX232	负调控抗旱 Negative regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	降低H₂O₂含量 Decreased H₂O₂content
MeGRX15	负调控抗旱 Negative regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	降低MDA含量 Decreased MDA content	[34]
MeGRX3	负调控抗旱 Negative regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	介导H₂O₂分配，引起ABA途径介导的气孔关闭 Mediated H₂O₂homeostasis and stomatal closure	[35]
MeCAT1 （Co-expressed with MeCu/ZnSOD）	正调控抗旱 Positive regulation of drought resistance	过氧化氢酶 Catalase	强化ROS清除 Enhanced ROS scavenging	[36]
	正调控抗寒 Positive regulation of cold resistance	超氧化物歧化酶 Superoxide dismutase	强化ROS清除 Enhanced ROS scavenging
MeMYB2	负调控抗旱 Negative regulation of drought resistance	MYB转录因子 MYB transcriptional factor	调节ABA依赖途径 Regulated ABA-dependent pathway	[37]
MeMYB2	负调控抗寒 Negative regulation of cold resistance	MYB转录因子 MYB transcriptional factor	调节ABA依赖途径 Regulated ABA-dependent pathway	[37]
MeNCED1/5	正调控抗旱 Positive regulation of drought resistance	ABA合成酶 ABA synthetase	促进ABA积累 Promoted ABA accumulation	[38]
MeCIPK23	正调控抗旱 Positive regulation of drought resistance	蛋白激酶 CBL interact protein kinase	促进ABA积累 Promoted ABA accumulation
MeWHY1/2/3	正调控抗旱 Positive regulation of drought resistance	Whirly转录因子 Whirly transcriptional factor	促进ABA积累 Promoted ABA accumulation
MeWRKY20	正调控抗旱 Positive regulation of drought resistance	WRKY转录因子 WRKY transcriptional factor	促进ABA积累 Promoted ABA accumulation	[39]
MeHSP90.9	正调控抗旱 Positive regulation of drought resistance	热激蛋白 Heat shock protein	and decrease H₂O₂content	[39]
MeSPL9	负调控抗旱 Negative regulation of drought resistance	SPL转录因子 SPL transcriptional factor	抑制花青素、脯氨酸、可溶性糖和JA积累 Inhibited anthocyanin, proline, soluble sugar, and JA accumulation	[40]
MeSDD1	正调控抗旱 Positive regulation of drought resistance	枯草杆菌蛋白酶 Subtilase	降低气孔密度 Decreased stomatal density	[41]
MeSCL30	正调控抗旱 Positive regulation of drought resistance	剪切因子 Spliceosomal component-like	强化ROS清除 Enhanced ROS scavenging	[42]
MeRSZ21b	正调控抗旱 Positive regulation of drought resistance	丝氨酸/精氨酸剪接因子 Two-Zn-knuckles-type serine/arginine-rich protein	调节ABA依赖途径的气孔关闭 Regulated stomatal closure of ABA-dependent pathway	[43]
DIR	正调控抗旱 Positive regulation of drought resistance	长链非编码RNA Long non-coding RNA	增加脯氨酸含量 Increased proline content	[44]
MeIPT	正调控抗旱 Positive regulation of drought resistance	异戊烯基转移酶 Isopentenyl transferase	延缓叶片衰老 Delayed leaf senescence	[45]
MeMYB108	正调控抗旱 Positive regulation of drought resistance	MYB转录因子 MYB transcriptional factor	降低叶片脱落率 Reduced leaf abscission rate	[46]
MeAPX2 （Co-expressed with MeCu/ZnSOD）	正调控抗寒 Positive regulation of cold resistance	抗坏血酸过氧化物酶 Ascorbate peroxidase	强化ROS清除 Enhanced ROS scavenging	[47]
MeTCP4	正调控抗寒 Positive regulation of drought resistance	TCP转录因子 TCP transcriptional factor	强化ROS清除 Enhanced ROS scavenging	[48]
CRIR1	正调控抗寒 Positive regulation of cold resistance	长链非编码RNA Long non-coding RNA	增强非CBF途径的低温胁迫相关基因的翻译 Improved the translation efficiency of cold stress-related genes in a CBF-independent pathway	[49]
MeNPF4.5	正调控氮利用效率和产量 Positive regulation of nitrogen use efficiency and yield	氮转运蛋白 Nitrate transporter	可能促进了IAA积累 May promote IAA accumulation	[50]
MeWRKY79	正调控CBB抗性 Positive regulation of CBB resistance	WRKY转录因子 WRKY transcriptional factor	促进褪黑素积累 Promoted melatonin accumulation	[51]
MeHsf20	正调控CBB抗性 Positive regulation of CBB resistance	热激蛋白 Heat shock protein	促进褪黑素积累 Promoted melatonin accumulation
MeASMT2	正调控CBB抗性 Positive regulation of CBB resistance	褪黑素合成酶 Melatonin synthetase	促进褪黑素积累 Promoted melatonin accumulation
MeHsfs3	正调控CBB抗性 Positive regulation of CBB resistance	热胁迫转录因子 Heat stress transcriptional factor	促进SA的积累 Promoted SA accumulation	[52]
MeDELLA1/2/3/4	正调控CBB抗性 Positive regulation of CBB resistance	DELLA蛋白 DELLA protein	促进胼胝质沉积 Promoted callose depostion	[53]
MeLRR1/2/3/4	正调控CBB抗性 Positive regulation of CBB resistance	NBS-LRR蛋白 NBS-LRR protein	促进SA和ROS积累 Promoted SA and ROS accumulation	[54]
MeWRKY27	正调控CBB抗性 Positive regulation of CBB resistance	WRKY转录因子 WRKY transcriptional factor	未知 Unknown	[55]
MeWRKY33	正调控CBB抗性 Positive regulation of CBB resistance	WRKY转录因子 WRKY transcriptional factor	未知 Unknown	[55]
MeWRKY20	正调控CBB抗性 Positive regulation of CBB resistance	WRKY转录因子 WRKY transcriptional factor	促进胼胝质增加，触发自噬信号途径 Promoted callose deposition, and triggered autophagy signaling pathway	[56]
MeHSP90.9	正调控CBB抗性 Positive regulation of CBB resistance	热激蛋白 Heat shock protein	触发自噬信号途径 Triggered autophagy signaling Pathway	[57]
MeDNAJA2	正调控CBB抗性 Positive regulation of CBB resistance	DnaJ热激蛋白 DnaJ heat shock protein family	促进SA积累 Promoted SA accumulation	[58]
MeHAM1	正调控CBB抗性 Positive regulation of CBB resistance	组蛋白乙酰转移 Histone acetyltransferases	促进SA积累 Promoted SA accumulation	[58]
MeLAR	正调控TMMS抗性 Positive regulation of TMMS resistance	无花色素还原酶 Leucoanthocyanidin reductase	促进单宁积累 Promoted tannin accumulation	[59]
MeANR	正调控TMMS抗性 Positive regulation of TMMS resistance	花青素还原酶Anthocyanin reductase	促进单宁积累 Promoted tannin accumulation	[59]

基因名称 Gene name	功能 Function	产物 Product	生理机制 Physiological mechanism	参考文献 Reference
MeCWINV3	负调控淀粉积累 Negative regulation of starch accumulation	细胞壁转化酶 Cell wall invertases	抑制叶片中蔗糖向块根转移Inhibited sugar export from leaves to storage roots	[25]
MeSBE1	负调控直链淀粉形成 Negative regulation of high-amylose production	淀粉分支酶1 Starch branch enzyme 1	促进块根中支链淀粉链长分布 Promoted chain-length distribution of amylopectin	[26]
MeSBE2	负调控直链淀粉形成 Negative regulation of high-amylose production	淀粉分支酶2 Starch branch enzyme 2	促进块根中支链淀粉链长分布 Promoted chain-length distribution of amylopectin	[26]
MeGBSSI	正调控糯木薯形成 Positive regulation of waxy cassava	颗粒结合淀粉合酶I Granule-bound starch synthase I	合成蜡质淀粉 Synthesized waxy starch	[27]
MePSY2	正调控黄色薯肉形成 Positive regulation of yellow tuber	八氢番茄红素酶2 Phytoene synthase 2	促进β-胡萝卜素合成 Promoted β-carotene accumulation	[14]
MeCAT1 （Co-expressed with MeCu/ZnSOD）	延缓PPD Delay PPD	过氧化氢酶 Catalase	强化ROS清除 Enhanced ROS scavenging	[28]
HNL	负调控块根中HCN含量 Negative regulation of HCN content in tuber	羟基腈裂解酶Hydroxynitrile lyase	促进丙酮氰醇分解 Promoted the decomposition of acetone cyanol	[29]
MeDREB1D	正调控抗旱 Positive regulation of drought resistance	AP2转录因子 AP2 transcriptional factor	降低MDA含量，并可能强化了ROS清除 Decreased MDA content, and may enhance ROS scavenging	[30]
MeDREB1D	正调控抗寒 Positive regulation of cold resistance	AP2转录因子 AP2 transcriptional factor	可能强化了ROS清除 May enhance ROS scavenging	[30]
MeDREB1A	正调控抗旱 Positive regulation of drought resistance	AP2转录因子 AP2 transcriptional factor	增加脯氨酸含量 Increased proline content	[31]
MeDREB1A	正调控抗寒 Positive regulation of cold resistance	AP2转录因子 AP2 transcriptional factor	增加脯氨酸含量 Increased proline content	[31]
MeRAV5	正调控抗旱 Positive regulation of drought resistance	AP2转录因子 AP2 transcriptional factor	降低H₂O₂含量和促进木质素积累 Decreased H₂O₂ content and promote lignin accumulation	[32]
MeGRX360	正调控抗旱 Positive regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	降低H₂O₂含量 Decreased H₂O₂content	[33]
MeGRX058	正调控抗旱 Positive regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	降低H₂O₂含量 Decreased H₂O₂content
MeGRX785	负调控抗旱 Negative regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	降低H₂O₂含量 Decreased H₂O₂content
MeGRX232	负调控抗旱 Negative regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	降低H₂O₂含量 Decreased H₂O₂content
MeGRX15	负调控抗旱 Negative regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	降低MDA含量 Decreased MDA content	[34]
MeGRX3	负调控抗旱 Negative regulation of drought resistance	CC型谷氧还蛋白 CC-type Glutaredoxins	介导H₂O₂分配，引起ABA途径介导的气孔关闭 Mediated H₂O₂homeostasis and stomatal closure	[35]
MeCAT1 （Co-expressed with MeCu/ZnSOD）	正调控抗旱 Positive regulation of drought resistance	过氧化氢酶 Catalase	强化ROS清除 Enhanced ROS scavenging	[36]
	正调控抗寒 Positive regulation of cold resistance	超氧化物歧化酶 Superoxide dismutase	强化ROS清除 Enhanced ROS scavenging
MeMYB2	负调控抗旱 Negative regulation of drought resistance	MYB转录因子 MYB transcriptional factor	调节ABA依赖途径 Regulated ABA-dependent pathway	[37]
MeMYB2	负调控抗寒 Negative regulation of cold resistance	MYB转录因子 MYB transcriptional factor	调节ABA依赖途径 Regulated ABA-dependent pathway	[37]
MeNCED1/5	正调控抗旱 Positive regulation of drought resistance	ABA合成酶 ABA synthetase	促进ABA积累 Promoted ABA accumulation	[38]
MeCIPK23	正调控抗旱 Positive regulation of drought resistance	蛋白激酶 CBL interact protein kinase	促进ABA积累 Promoted ABA accumulation
MeWHY1/2/3	正调控抗旱 Positive regulation of drought resistance	Whirly转录因子 Whirly transcriptional factor	促进ABA积累 Promoted ABA accumulation
MeWRKY20	正调控抗旱 Positive regulation of drought resistance	WRKY转录因子 WRKY transcriptional factor	促进ABA积累 Promoted ABA accumulation	[39]
MeHSP90.9	正调控抗旱 Positive regulation of drought resistance	热激蛋白 Heat shock protein	and decrease H₂O₂content	[39]
MeSPL9	负调控抗旱 Negative regulation of drought resistance	SPL转录因子 SPL transcriptional factor	抑制花青素、脯氨酸、可溶性糖和JA积累 Inhibited anthocyanin, proline, soluble sugar, and JA accumulation	[40]
MeSDD1	正调控抗旱 Positive regulation of drought resistance	枯草杆菌蛋白酶 Subtilase	降低气孔密度 Decreased stomatal density	[41]
MeSCL30	正调控抗旱 Positive regulation of drought resistance	剪切因子 Spliceosomal component-like	强化ROS清除 Enhanced ROS scavenging	[42]
MeRSZ21b	正调控抗旱 Positive regulation of drought resistance	丝氨酸/精氨酸剪接因子 Two-Zn-knuckles-type serine/arginine-rich protein	调节ABA依赖途径的气孔关闭 Regulated stomatal closure of ABA-dependent pathway	[43]
DIR	正调控抗旱 Positive regulation of drought resistance	长链非编码RNA Long non-coding RNA	增加脯氨酸含量 Increased proline content	[44]
MeIPT	正调控抗旱 Positive regulation of drought resistance	异戊烯基转移酶 Isopentenyl transferase	延缓叶片衰老 Delayed leaf senescence	[45]
MeMYB108	正调控抗旱 Positive regulation of drought resistance	MYB转录因子 MYB transcriptional factor	降低叶片脱落率 Reduced leaf abscission rate	[46]
MeAPX2 （Co-expressed with MeCu/ZnSOD）	正调控抗寒 Positive regulation of cold resistance	抗坏血酸过氧化物酶 Ascorbate peroxidase	强化ROS清除 Enhanced ROS scavenging	[47]
MeTCP4	正调控抗寒 Positive regulation of drought resistance	TCP转录因子 TCP transcriptional factor	强化ROS清除 Enhanced ROS scavenging	[48]
CRIR1	正调控抗寒 Positive regulation of cold resistance	长链非编码RNA Long non-coding RNA	增强非CBF途径的低温胁迫相关基因的翻译 Improved the translation efficiency of cold stress-related genes in a CBF-independent pathway	[49]
MeNPF4.5	正调控氮利用效率和产量 Positive regulation of nitrogen use efficiency and yield	氮转运蛋白 Nitrate transporter	可能促进了IAA积累 May promote IAA accumulation	[50]
MeWRKY79	正调控CBB抗性 Positive regulation of CBB resistance	WRKY转录因子 WRKY transcriptional factor	促进褪黑素积累 Promoted melatonin accumulation	[51]
MeHsf20	正调控CBB抗性 Positive regulation of CBB resistance	热激蛋白 Heat shock protein	促进褪黑素积累 Promoted melatonin accumulation
MeASMT2	正调控CBB抗性 Positive regulation of CBB resistance	褪黑素合成酶 Melatonin synthetase	促进褪黑素积累 Promoted melatonin accumulation
MeHsfs3	正调控CBB抗性 Positive regulation of CBB resistance	热胁迫转录因子 Heat stress transcriptional factor	促进SA的积累 Promoted SA accumulation	[52]
MeDELLA1/2/3/4	正调控CBB抗性 Positive regulation of CBB resistance	DELLA蛋白 DELLA protein	促进胼胝质沉积 Promoted callose depostion	[53]
MeLRR1/2/3/4	正调控CBB抗性 Positive regulation of CBB resistance	NBS-LRR蛋白 NBS-LRR protein	促进SA和ROS积累 Promoted SA and ROS accumulation	[54]
MeWRKY27	正调控CBB抗性 Positive regulation of CBB resistance	WRKY转录因子 WRKY transcriptional factor	未知 Unknown	[55]
MeWRKY33	正调控CBB抗性 Positive regulation of CBB resistance	WRKY转录因子 WRKY transcriptional factor	未知 Unknown	[55]
MeWRKY20	正调控CBB抗性 Positive regulation of CBB resistance	WRKY转录因子 WRKY transcriptional factor	促进胼胝质增加，触发自噬信号途径 Promoted callose deposition, and triggered autophagy signaling pathway	[56]
MeHSP90.9	正调控CBB抗性 Positive regulation of CBB resistance	热激蛋白 Heat shock protein	触发自噬信号途径 Triggered autophagy signaling Pathway	[57]
MeDNAJA2	正调控CBB抗性 Positive regulation of CBB resistance	DnaJ热激蛋白 DnaJ heat shock protein family	促进SA积累 Promoted SA accumulation	[58]
MeHAM1	正调控CBB抗性 Positive regulation of CBB resistance	组蛋白乙酰转移 Histone acetyltransferases	促进SA积累 Promoted SA accumulation	[58]
MeLAR	正调控TMMS抗性 Positive regulation of TMMS resistance	无花色素还原酶 Leucoanthocyanidin reductase	促进单宁积累 Promoted tannin accumulation	[59]
MeANR	正调控TMMS抗性 Positive regulation of TMMS resistance	花青素还原酶Anthocyanin reductase	促进单宁积累 Promoted tannin accumulation	[59]