主要麻类作物基因组学与遗传改良: 现状与展望

doi:10.3724/SP.J.1006.2021.04121

作物学报 ›› 2021, Vol. 47 ›› Issue (6): 997-1019.doi: 10.3724/SP.J.1006.2021.04121

• 专题：主要麻类作物基因组学与遗传改良 • 上一篇下一篇

主要麻类作物基因组学与遗传改良: 现状与展望

徐益¹^,²^,³(), 张力岚¹^,²^,³, 祁建民¹^,², 张列梅¹, 张立武¹^,²^,³^,^*()

¹福建农林大学农学院/作物遗传育种与综合利用教育部重点实验室/福建省作物设计育种重点实验室, 福建福州 350002
²福建农林大学农业农村部东南黄红麻实验观测站/福建省麻类种质资源共享平台/福建省南方经济作物遗传育种与多用途开发国际科技合作基地, 福建福州 350002
³福建农林大学海峡联合研究院基因组与生物技术中心, 福建福州 350002

收稿日期:2020-05-01 接受日期:2021-01-11 出版日期:2021-06-12 网络出版日期:2021-01-25
通讯作者: 张立武
作者简介:E-mail:1275924118@qq.com
基金资助:
国家自然科学基金项目(31771369);国家自然科学基金项目(31972968);国家现代农业产业技术体系建设专项(CARS-19-E06)

Genomics and genetic improvement in main bast fiber crops: advances and perspectives

XU Yi¹^,²^,³(), ZHANG Li-Lan¹^,²^,³, QI Jian-Min¹^,², ZHANG Lie-Mei¹, ZHANG Li-Wu¹^,²^,³^,^*()

¹Key Laboratory of Ministry of Education for Genetics, Breeding and Multiple Utilization of Crops/Fujian Key Laboratory for Crop Breeding by Design/College of Agriculture, Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
²Experiment Station of Jute and Kenaf in Southeast China, Ministry of Agriculture and Rural Affairs/Public Platform of Fujian for Germplasm Resources of Bast Fiber Crops/Fujian International Science and Technology Cooperation Base for Genetics, Breeding and Multiple Utilization Development of Southern Economic Crops, Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
³Center for Genomics and Biotechnology, Haixia Institute of Science and Technology, Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China

Received:2020-05-01 Accepted:2021-01-11 Published:2021-06-12 Published online:2021-01-25
Contact: ZHANG Li-Wu
Supported by:
The National Natural Science Foundation of China(31771369);The National Natural Science Foundation of China(31972968);The China Agriculture Research System(CARS-19-E06)

摘要/Abstract

摘要：

随着测序技术的发展, 主要麻类作物(黄麻、红麻、苎麻、亚麻和工业大麻)参考基因组从2011年至2020年陆续完成测序, 这标志着麻类作物科学已经进入基因组时代。文章首先详细概述主要麻类作物基因组测序。其次, 评述了基于基因组学的麻类作物重要应用价值基因挖掘。基于参考基因组和转录组测序, 大量关于纤维发育、响应非生物胁迫的候选基因被挖掘, 以促进麻类作物纤维的物种特性和“不与粮食争好地”的逆境农业。同时不同麻类作物特异性状候选基因陆续被报道, 如红麻雄性不育、亚麻种子含油量和大麻大麻素相关候选基因等。再次, 麻类作物基因组测序完成为基于组学的麻类作物遗传改良提供可能: 有助于麻类作物种质资源形成和演化机制研究, 系统解析纤维产量、纤维品质、抗病耐逆等农艺性状形成的分子基础; 有助于建立高通量基因型-表型数据库, 挖掘优异基因资源与创制新种质; 有助于创新并集成分子标记辅助选择、基因组选择、转基因等技术, 建立高效的快速育种技术体系。宜选育高产高效、抗逆抗病、适宜轻简化机械化、优质专用的多用途麻类作物新品种, 以满足麻类作物相关产业的市场需求, 适应麻类作物生产方式。尽管已经获得重要基因以及位点的信息, 但如何高效率利用已有基因资源对麻类作物进行遗传改良仍需面临一系列挑战, 如成熟稳定的遗传转化体系、麻类作物基因编辑体系构建及基因组选择育种等。

关键词: 主要麻类作物, 基因组, 基因, 遗传改良

Abstract:

With the development of sequencing technology, main bast fiber crops (jute, kenaf, ramie, flax, and hemp) have completed genome sequencing from 2011 to 2020, which marks that the science of bast fiber crops has entered the era of genome. Firstly, this paper reviews the genome sequencing of bast fiber crops. Secondly, the important gene identification of bast fiber crops is also reported. Based on reference genome and transcriptome sequencing, a large number of candidate genes related to fiber development and response to abiotic stress have been detected, corresponding to the species characteristics of bast fiber and the adversity agriculture of “not competing with food”. Meanwhile, candidate genes for specific bast fiber crops have also been identified, such as male fertility in kenaf, seed oil content in flax, cannabinoid related candidate genes. Thirdly, the completion of bast fiber crop genome sequencing provides the possibility of omics-based genetic improvement, which will facilitate to study the formation of bast fiber and evolution mechanisms of bast fiber crop germplasms and systematically analyze the molecular basis for the formation of agronomic traits such as fiber yield, fiber quality, disease resistance, and stress tolerance. Also, it will facilitate to establish a high-throughput genotype-phenotype database, mine excellent gene resources, and create new germplasm. Moreover, it will facilitate to establish efficient rapid breeding technology systems by the innovation and combination of molecular marker-assisted selection, genome selection, transgenic technology and so on. To meet the market demand particularly bast fiber crop-related industries and adapt to the production model of bast fiber crops, we should breed new bast fiber crop varieties with high yield, high efficiency, stress resistance, disease resistance, suitable for light simplification and mechanization cultivation, high quality, and special purpose. Although the important information of gene resources and loci has been obtained from the reference genomes, there are still a series of challenges that how to utilize the existing resources efficiently for genetic improvement of bast fiber crops, such as stable and efficient genetic transformation system, construction of gene editing system, and genome selection breeding.

Key words: major bast fiber crops, genome, gene, genetic improvement

徐益, 张力岚, 祁建民, 张列梅, 张立武. 主要麻类作物基因组学与遗传改良: 现状与展望[J]. 作物学报, 2021, 47(6): 997-1019.

XU Yi, ZHANG Li-Lan, QI Jian-Min, ZHANG Lie-Mei, ZHANG Li-Wu. Genomics and genetic improvement in main bast fiber crops: advances and perspectives[J]. Acta Agronomica Sinica, 2021, 47(6): 997-1019.

图/表 6

图1

表1

附图1

附表1

主要麻类作物纤维发育候选基因信息"

基因 Gene	作物 Crop					功能或表型 Function or phenotype								文献 Reference
4CL	黄麻, 红麻 Jute, kenaf					参与木质素生物合成 Involved in lignin biosynthesis								[6, 8, 20]
4CL1	苎麻 Ramie					参与木质素生物合成 Involved in lignin biosynthesis								[47]
4CL3	苎麻 Ramie					参与黄酮类化合物的合成 Involved in the synthesis of flavonoids								[47]
APL	黄麻, 红麻 Jute, kenaf					参与纤维素生物合成 Involved in cellulose biosynthesis								[6, 8]
bHLH	亚麻 Flax					参与茎底部区域次生细胞壁沉积 Involved in the deposition of secondary cell wall at the bottom of stem								[66]
基因 Gene		作物 Crop					功能或表型 Function or phenotype					文献 Reference
bZIP		亚麻 Flax					参与茎底部区域次生细胞壁沉积 Involved in the deposition of secondary cell wall at the bottom of stem						[66]
C3H		红麻 Kenaf					参与木质素生物合成 Involved in lignin biosynthesis						[24]
C4H		红麻, 苎麻 Kenaf, ramie					参与木质素生物合成 Involved in lignin biosynthesis						[8, 47]
CAD		黄麻, 红麻, 苎麻 Jute, kenaf, ramie					参与纤维素生物合成 Involved in cellulose biosynthesis						[19, 24, 47]
CCR		黄麻, 红麻 Jute, kenaf					参与木质素生物合成 Involved in lignin biosynthesis						[6, 8, 20]
CCoAOMT		黄麻,红麻,苎麻 Jute, kenaf, ramie					参与木质素生物合成 Involved in lignin biosynthesis						[6, 24, 47]
CesA		亚麻 Flax					沉默后, 韧皮纤维的数量和结构受到了严重的影响 The quantity and structure of phloem fibers were seriously affected after silencing						[62-63]
CesA1		红麻, 苎麻 Kenaf, ramie					参与初生细胞壁的纤维素沉积 Involved in cellulose deposition of primary cell wall						[8, 42]
CesA2		苎麻 Ramie					参与木质素生物合成 Involved in lignin biosynthesis						[43]
CesA3		红麻, 苎麻 Kenaf, ramie					参与初生细胞壁的纤维素沉积 Involved in cellulose deposition of primary cell wall						[8, 43]
CesA4		黄麻, 红麻, 苎麻, 工业大麻 Jute, kenaf, ramie, hemp					参与次生细胞壁中纤维素沉积 Involved in cellulose deposition in secondary cell wall						[6, 8, 44, 80]
CesA6		红麻 Kenaf					参与初生细胞壁的纤维素沉积 Involved in cellulose deposition of primary cell wall						[8]
CesA7		黄麻, 红麻, 工业大麻 Jute, kenaf, hemp					协调木聚糖型S层的沉积 Coordinating the S-layers deposition in the xylan-type						[8, 19, 80]
CesA8		红麻, 工业大麻 Kenaf, hemp					参与次生细胞壁中纤维素沉积 Involved in cellulose deposition in secondary cell wall						[8, 80]
CML15b		亚麻 Flax					PLR1基因转录调控的关键因子 Key factors for transcription regulation of PLR1 gene						[68]
COB		工业大麻 Hemp					在次级生长的下胚轴中有较高表达量 Highly expressed in hypocotyls undergoing secondary growth						[80]
COBL4		工业大麻 Hemp					在次级生长的下胚轴中有较高表达量 Highly expressed in hypocotyls undergoing secondary growth						[80]
COMT		黄麻, 红麻, 亚麻 Jute, kenaf, flax					参与木质素生物合成 Involved in lignin biosynthesis						[6, 8, 20, 24, 67]
CSL		黄麻, 红麻, 亚麻 Jute, kenaf, flax					参与纤维素生物合成 Involved in cellulose biosynthesis						[8, 17, 62]
F5H		红麻, 苎麻 Kenaf, ramie					参与木质素生物合成 Involved in lignin biosynthesis						[8, 24, 47]
FLA6		黄麻 Jute					协调木聚糖型S层的沉积 Coordinating the S-layers deposition in the xylan-type						[19]
FLA11		工业大麻 Hemp					影响植物细胞壁中纤维素、阿拉伯糖和半乳糖的含量 Impact cellulose, arabinose and galactose content in plant cell walls						[80, 82]
FLA12		工业大麻 Hemp					影响植物细胞壁中纤维素、阿拉伯糖和半乳糖的含量 Impact cellulose, arabinose and galactose content in plant cell walls						[80, 82]
G2-like		亚麻 Flax					参与茎底部区域次生细胞壁沉积 Involved in the deposition of secondary cell wall at the bottom of stem						[66]
GRAS		亚麻 Flax					分支数候选基因Candidate gene for the number of branches						[112]
基因 Gene			作物 Crop					功能或表型 Function or phenotype		文献 Reference
HAT22			黄麻, 红麻 Jute, kenaf					参与纤维素生物合成 Involved in cellulose biosynthesis			[6, 8]
HB8			红麻 Kenaf					参与纤维形成 Involved in fiber formation			[8]
HCA2			红麻 Kenaf					参与纤维形成 Involved in fiber formation			[8]
HCT			红麻 Kenaf					参与木质素生物合成 Involved in lignin biosynthesis			[8, 24]
KANADI			红麻 Kenaf					参与纤维形成 Involved in fiber formation			[8]
Kor			黄麻, 红麻 Jute, kenaf					参与纤维素生物合成 Involved in cellulose biosynthesis			[8, 17]
LBD1			红麻 Kenaf					参与纤维形成 Involved in fiber formation			[8]
MP			红麻 Kenaf					参与纤维形成 Involved in fiber formation			[8]
MYB			亚麻 Flax					参与茎底部区域次生细胞壁沉积 Involved in the deposition of secondary cell wall at the bottom of stem			[66]
MYB46			红麻, 苎麻, 亚麻, 工业大麻 Kenaf, ramie, flax, hemp					参与次生细胞壁生物发生和木聚糖生物合成 Involved in secondary cell wall biogenesis and xylan biosynthesis			[8, 12, 48, 80, 82]
MYB58			红麻 Kenaf					参与韧皮纤维的次生细胞壁合成 Involved in the secondary cell wall synthesis of phloem fibers			[8]
MYB83			黄麻, 红麻, 亚麻 Jute, kenaf, flax					参与纤维素生物合成 Involved in cellulose biosynthesis			[6, 8, 12]
MYB85			红麻 Kenaf					参与韧皮纤维的次生细胞壁合成 Involved in the secondary cell wall synthesis of phloem fibers			[8]
MYB103			红麻 Kenaf					参与韧皮纤维的次生细胞壁合成 Involved in the secondary cell wall synthesis of phloem fibers			[8]
NAC19			苎麻 Ramie					上调MYB46的表达 Up regulation of MYB46 expression			[48]
NAC24			苎麻 Ramie					上调MYB46的表达 Up regulation of MYB46 expression			[48]
NST1			工业大麻 Hemp					纤维分化的主要调节因子 Major regulator of fibre differentiation			[80-81]
PAL			红麻, 苎麻 Kenaf, ramie					参与木质素生物合成 Involved in lignin biosynthesis			[8, 47]
NST1			红麻 Kenaf					参与韧皮纤维的次生细胞壁合成 Involved in the secondary cell wall synthesis of phloem fibers			[8]
PHAC1			亚麻 Flax					表达量与纤维拉力强度成正比 The expression is proportional to the tensile strength of the fiber			[68]
PIN1			红麻 Kenaf					参与纤维形成 Involved in fiber formation			[8]
PL			亚麻 Flax					株高候选基因 Candidate gene for plant height			[112]
PLR1			亚麻 Flax					参与木质素生物合成 Involved in lignin biosynthesis			[68]
ROPGAP3			亚麻 Flax					参与韧皮纤维的次生细胞壁合成 Involved in the secondary cell wall synthesis of phloem fibers			[12]
SND1			红麻 Kenaf					参与韧皮纤维的次生细胞壁合成 Involved in the secondary cell wall synthesis of phloem fibers			[8]
SND2			红麻 Kenaf					参与韧皮纤维的次生细胞壁合成 Involved in the secondary cell wall synthesis of phloem fibers			[8]
SND3			红麻 Kenaf					参与韧皮纤维的次生细胞壁合成 Involved in the secondary cell wall synthesis of phloem fibers			[8]
Susy			黄麻, 红麻 Jute, kenaf					参与纤维素生物合成 Involved in cellulose biosynthesis			[8, 17]
基因 Gene				作物 Crop	功能或表型 Function or phenotype				文献 Reference
TDIF				红麻 Kenaf	参与纤维形成 Involved in fiber formation				[8]
TOUCH4				工业大麻 Hemp	将木葡聚糖靶向于次生细胞壁S1层的初生-次生细胞壁交界 Participate in the targeting of xyloglucan to the primary- secondary cell wall junction in the secondary cell wall S1 layer				[80]
UGPase				黄麻 Jute	参与纤维素生物合成 Involved in cellulose biosynthesis				[17]
UGT				亚麻 Flax	株高候选基因 Candidate gene for plant height				[112]
WAT1				苎麻 Ramie	高纤维品种驯化中的受选择基因 Selection of highly domesticated fiber varieties				[45]
WLIM1				工业大麻 Hemp	捆绑肌动蛋白丝来促进纤维的延伸, 与PAL-box结合来促进木质素生物合成基因的木质化 Fibre extension by bundling the actin filament but also fibre lignification by promoting the lignin/lignin-like biosynthetic genes via binding to the PAL-box				[80, 83]
WOX4				黄麻, 红麻 Jute, kenaf	参与纤维素生物合成 Involved in cellulose biosynthesis				[6, 8]
WRKY				黄麻 Jute	纤维改良 Fiber improvement				[20]
WRKY36				亚麻 Flax	通过与PLR1启动子中的W-box结合, 参与木质素生物合成 Involved in lignin biosynthesis by binding with W-box in PLR1 promoter				[69]
XTH				亚麻 Flax	分支数候选基因 Candidate gene for the number of branches				[112]
XTH5				工业大麻 Hemp	韧皮纤维的伸长依赖于XTH的活性 Bast fibre extension depends on the activities of XTH				[80]
XTH8				工业大麻 Hemp	韧皮纤维的伸长依赖于XTH的活性 Bast fibre extension depends on the activities of XTH				[80]
XTH15				工业大麻 Hemp	在二次生长的较老大麻下胚轴中表达较多 More expressed in the elongating hemp hypocotyl				[80]
XTH22				工业大麻 Hemp	在二次生长的较老大麻下胚轴中表达较多 More expressed in the elongating hemp hypocotyl				[80]

附表1

附表2

主要麻类作物响应非生物胁迫候选基因信息"

基因 Gene	作物 Crop				功能或表型 Function or phenotype					文献 Reference
α-amylase	苎麻 Ramie				响应干旱和高盐逆境胁迫 Response to drought and high salt stress					[53]
A-ARR	黄麻 Jute				通过细胞分裂素途径响应盐胁迫 Response to salt stress through cytokinin pathway					[21]
ABF	黄麻 Jute				通过ABA信号通路响应盐胁迫 Response to salt stress through the ABA signaling pathway					[21]
ACO1	苎麻 Ramie				响应干旱和高盐逆境胁迫 Response to drought and high salt stress					[51]
AHP	黄麻 Jute				通过细胞分裂素途径响应盐胁迫 Response to salt stress through cytokinin pathway					[21]
AP2/ERF	红麻 Kenaf				在盐胁迫转录组中检测到的差异表达因子 Differential expression factors detected in transcriptome under salt stress					[27]
AP2/EREBP	红麻 Kenaf				在盐胁迫转录组中检测到的差异表达因子 Differential expression factors detected in transcriptome under salt stress					[27]
ARF	红麻 Kenaf				在盐胁迫转录组中检测到的差异表达因子 Differential expression factors detected in transcriptome under salt stress					[27]
基因 Gene			作物 Crop			功能或表型 Function or phenotype			文献 Reference
AvrL2		亚麻 Flax				抵御锈病感染 Resist rust infection			[76, 107]
AvrL567		亚麻 Flax				抵御锈病感染 Resist rust infection			[76]
AvrM14		亚麻 Flax				抵御锈病感染 Resist rust infection			[76, 107]
AvrP123		亚麻 Flax				抵御锈病感染 Resist rust infection			[76]
AvrP4		亚麻 Flax				抵御锈病感染 Resist rust infection			[76]
AvrM		亚麻 Flax				抵御锈病感染 Resist rust infection			[76]
B-ARR		黄麻 Jute				通过细胞分裂素途径响应盐胁迫 Response to salt stress through cytokinin pathway			[21]
bHLH		红麻 Kenaf				在盐胁迫转录组中检测到的差异表达因子 Differential expression factors detected in transcriptome under salt stress			[27]
bZIP		红麻, 苎麻 Kenaf, ramie				响应干旱和高盐逆境胁迫 Response to drought and high salt stress			[27, 52]
CAX3		亚麻 Flax				基因产物可能通过Ca²⁺介导的胞内调节参与了亚麻对高酸性、高Al³⁺浓度的响应 Gene product may be involved in the response of flax to high acidity and high Al³⁺ concentration through Ca²⁺ mediated intracellular regulation			[73]
CCAAT		黄麻 Jute				在盐胁迫转录组中检测到的差异表达因子 Differential expression factors detected in transcriptome under salt stress			[21]
CRE1		黄麻 Jute				通过细胞分裂素途径响应盐胁迫 Response to salt stress through cytokinin pathway			[21]
G6PDH1		苎麻 Ramie				响应重金属镉胁迫 Response to cadmium stress			[57]
GS2		苎麻 Ramie				转基因烟草能提高生物产量和氮利用效率 Transgenic tobacco can improve biomass and nitrogen utilization			[54]
HB		黄麻 Jute				在盐胁迫转录组中检测到的差异表达因子 Differential expression factors detected in transcriptome under salt stress			[21]
HDA2		红麻 Kenaf				盐胁迫和干旱胁迫的响应基因 Response genes of salt and drought stress			[30]
HDA8		红麻 Kenaf				盐胁迫和干旱胁迫的响应基因 Response genes of salt and drought stress			[30]
HDA9		红麻 Kenaf				盐胁迫和干旱胁迫的响应基因 Response genes of salt and drought stress			[30]
HDA19		红麻 Kenaf				盐胁迫和干旱胁迫的响应基因 Response genes of salt and drought stress			[30]
HHDA6		红麻 Kenaf				盐胁迫和干旱胁迫的响应基因 Response genes of salt and drought stress			[30]
HSF		黄麻 Jute				在盐胁迫转录组中检测到的差异表达因子 Differential expression factors detected in transcriptome under salt stress			[21]
HSF		亚麻 Flax				响应高温胁迫 Response to high temperature stress			[70]
JAS		亚麻 Flax				响应土壤营养胁迫的响应 Response to soil nutrient stress			[71]
KCS		黄麻 Jute				转基因增强了植株的抗旱性 Transgene enhances plant drought resistance			[23]
MADS-box		亚麻 Flax				参与调节植物生长发育和参与耐铝性细胞壁修饰的酶 Enzymes involved in the regulation of plant growth and development and in the modification of aluminum resistant cell wall			[74]
MYB		黄麻 Jute				在盐胁迫转录组中检测到的差异表达因子 Differential expression factors detected in transcriptome under salt stress			[21]
MYB83		苎麻 Ramie				响应重金属镉胁迫 Response to cadmium stress			[56]
NAC		红麻 Kenaf				在盐胁迫转录组中检测到的差异表达因子 Differential expression factors detected in transcriptome under salt stress			[27]
基因 Gene				作物 Crop			功能或表型 Function or phenotype	文献 Reference
NAC				亚麻 Flax			参与调节植物生长发育和参与耐铝性细胞壁修饰的酶 Enzymes involved in the regulation of plant growth and development and in the modification of aluminum resistant cell wall	[74]
NRAMP1				苎麻 Ramie			响应重金属镉胁迫 Response to cadmium stress	[58]
PCS1				苎麻 Ramie			响应重金属镉胁迫 Response to cadmium stress	[55]
PP2C				黄麻 Jute			通过ABA信号通路响应盐胁迫 Response to salt stress through the ABA signaling pathway	[21]
PYL				黄麻 Jute			通过ABA信号通路响应盐胁迫 Response to salt stress through the ABA signaling pathway	[21]
RLK				黄麻 Jute			抗旱机制的负调控因子 Negative regulatory factors of drought resistance mechanism	[22]
SnRK2				黄麻 Jute			通过ABA信号通路响应盐胁迫 Response to salt stress through the ABA signaling pathway	[21]
SRT2				红麻 Kenaf			盐胁迫和干旱胁迫的响应基因 Response genes of salt and drought stress	[30]
TCP				红麻 Kenaf			在盐胁迫转录组中检测到的差异表达因子 Differential expression factors detected in transcriptome under salt stress	[27]
WD40-1				红麻 Kenaf			ABA和MeJA信号转导途径、盐和干旱胁迫应答途径的关键枢纽基因 Key genes of ABA and MeJA signal transduction pathway, salt and drought stress response pathway	[28]
WRKY				黄麻, 红麻, 苎麻 Jute, kenaf, ramie			在盐胁迫和干旱胁迫转录组中检测到的差异表达因子 Differential expression factors detected in transcriptome under salt and drought stress	[21, 27, 29, 50]

附表2

附表3

主要麻类作物特异性状候选基因信息"

基因 Gene		作物 Crop		功能或表型 Function or phenotype		文献 Reference
LMI1		红麻 Kenaf		叶形基因 Leaf shape gene		[8]
GI		红麻 Kenaf		参与红麻光周期调控 Involved in the photoperiod regulation of kenaf		[26]
CO		红麻 Kenaf		参与红麻光周期调控 Involved in the photoperiod regulation of kenaf		[26]
MADS-box		红麻 Kenaf		可能参与了红麻细胞质雄性不育 May be involved in the cytoplasmic male sterility of kenaf		[35]
TIR1		红麻 Kenaf		编码一个富含亮氨酸重复的F-box蛋白 Encodes a leucine rich F-box protein		[38-39]
atp9		红麻, 苎麻 Kenaf, ramie		细胞质雄性不育 Cytoplasmic male sterility		[32, 61]
atp6		红麻, 苎麻 Kenaf, ramie		细胞质雄性不育 Cytoplasmic male sterility		[35-37, 61]
ATPA		苎麻 Ramie		育性相关的基因 Fertility related genes		[60]
PLA2		亚麻 Flax		参与甘油磷脂代谢途径和醚酯代谢途径 Involved in glycerophosphatidic and ether ester metabolic pathways		[77]
PLC		亚麻 Flax		参与甘油磷脂代谢和醚酯代谢和肌醇磷酸代谢 Involved in the metabolism of glycerophosphatide, ether ester and inositol phosphate		[77]
PDAT1		亚麻 Flax		参与TAG合成途径 Involved in TAG synthesis pathway		[78]
DGAT1		亚麻 Flax		参与TAG合成途径 Involved in TAG synthesis pathway		[78]
DGAT2		亚麻 Flax		参与TAG合成途径 Involved in TAG synthesis pathway		[78]
FAD2a		亚麻 Flax		参与不饱和脂肪酸积累 Involved in the accumulation of unsaturated fatty acids		[79]
FAD3a		亚麻 Flax		参与不饱和脂肪酸积累 Involved in the accumulation of unsaturated fatty acids		[79]
基因 Gene	作物 Crop		功能或表型 Function or phenotype		文献 Reference
FAD3b	亚麻 Flax		参与不饱和脂肪酸积累 Involved in the accumulation of unsaturated fatty acids		[79]
PHO1	亚麻 Flax		千粒重候选基因Candidate gene for the 1000-seed weight		[112]
AEE1	工业大麻 Hemp		参与调控大麻素合成途径 Involved in the regulation of cannabinoid synthesis pathway		[13]
OLS	工业大麻 Hemp		参与调控大麻素合成途径 Involved in the regulation of cannabinoid synthesis pathway		[13, 86]
THCAS-like1	工业大麻 Hemp		编码大麻素原酸形成 Encodes cannabinoid acid formation		[13]
THCAS-like2	工业大麻 Hemp		编码大麻素原酸形成 Encodes cannabinoid acid formation		[13]
THCAS-like3	工业大麻 Hemp		编码大麻素原酸形成 Encodes cannabinoid acid formation		[13]
THCAS-like4	工业大麻 Hemp		编码大麻素原酸形成 Encodes cannabinoid acid formation		[13]
CMK	工业大麻 Hemp		表达量和大麻素含量呈现显著正相关 The expression level was positively correlated with cannabinoid content		[86]
MDS	工业大麻 Hemp		表达量和大麻素含量呈现显著正相关 The expression level was positively correlated with cannabinoid content		[86]
HDS	工业大麻 Hemp		表达量和大麻素含量呈现显著正相关 The expression level was positively correlated with cannabinoid content		[86]
HDR	工业大麻 Hemp		表达量和大麻素含量呈现显著正相关 The expression level was positively correlated with cannabinoid content		[86]
GPP(lsu)	工业大麻 Hemp		表达量和大麻素含量呈现显著正相关 The expression level was positively correlated with cannabinoid content		[86]
PT	工业大麻 Hemp		在始果期苞片腺毛中对大麻素合成积累起着关键作用 Played key roles in the biosynthesis and accumulation of cannabinoids in the glandular hairs of bract at initial-fruiting stage		[86]
THCAS	工业大麻 Hemp		在始果期苞片腺毛中对大麻素合成积累起着关键作用 Played key roles in the biosynthesis and accumulation of cannabinoids in the glandular hairs of bract at initial-fruiting stage		[86]
TPS18VF	工业大麻 Hemp		编码橙花醇/芳樟醇合成酶 Nerolidol/linalool synthases		[87]
TPS19BL	工业大麻 Hemp		编码橙花醇/芳樟醇合成酶 Nerolidol/linalool synthases		[87]
TPS16CC	工业大麻 Hemp		编码大根香叶烯B合酶 Germacrene B synthase		[87]
TPS20CT	工业大麻 Hemp		编码四甲基环癸二烯甲醇合酶 Hedycaryol synthase		[87]

附表3

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作物 Crop		测序技术 Sequencing technology	基因组大小 Genome size (Mb)	科 Families	染色体数目 Number of chromosomes	繁殖方式 Reproduction mode	文献 Reference
黄麻 Corchorus capsularis	圆果种 C. capsularis	二代测序 NGS	338	锦葵科 Malvaceae	2n=14	自花授粉 Self-pollination	[6]
	长果种 C. olitorius	二代测序 NGS	445			常异花授粉 Often cross-pollination	[6]
	圆果种 C. capsularis	三代测序 TGS	336				[7]
	长果种 C. olitorius	三代测序 TGS	361				[7]
红麻 Hibiscus cannabinus		二代+三代测序技术 NGS and TGS	1078	锦葵科 Malvaceae	2n=36	常异花授粉 Often cross-pollination	[8]
苎麻 Boehmeria nivea		二代测序 NGS	342	荨麻科 Urticaceae	2n=28	无性繁殖 Asexual reproduction/异花授粉Cross-pollination	[9]
苎麻 Boehmeria nivea		二代测序 NGS	336	荨麻科 Urticaceae	2n=28	无性繁殖 Asexual reproduction/异花授粉Cross-pollination	[10]
亚麻 Linum usitatissimum		二代测序 NGS	373	亚麻科 Linaceae	2n=30	自花授粉 Self-pollination	[11]
亚麻 Linum usitatissimum		二代测序 NGS	306,304,294	亚麻科 Linaceae		自花授粉 Self-pollination	[12]
工业大麻 Cannabis sativa		二代测序 NGS	534	大麻科 Cannabinaceae	2n=20	异花授粉 Cross-pollination	[13]
工业大麻 Cannabis sativa		二代测序 NGS	808	大麻科 Cannabinaceae	2n=20	异花授粉 Cross-pollination	[14]

主要麻类作物基因组学与遗传改良: 现状与展望

Genomics and genetic improvement in main bast fiber crops: advances and perspectives

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