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作物学报 ›› 2021, Vol. 47 ›› Issue (12): 2299-2313.doi: 10.3724/SP.J.1006.2021.14022

• 综述 •    下一篇

大豆种子颜色遗传调控机制研究进展

邱红梅1(), 陈亮1, 侯云龙1, 王新风1, 陈健1, 马晓萍1, 崔正果1, 张玲1, 胡金海1, 王跃强1,*(), 邱丽娟2,*()   

  1. 1吉林省农业科学院 / 大豆国家工程研究中心, 吉林长春 130033
    2中国农业科学院作物科学研究所 / 农作物基因资源与遗传改良国家重大科学工程, 北京 100081
  • 收稿日期:2021-01-24 接受日期:2021-05-17 出版日期:2021-12-12 网络出版日期:2021-05-26
  • 通讯作者: 王跃强,邱丽娟
  • 作者简介:E-mail: qhm2001-2005@163.com
  • 基金资助:
    “十三五”国家重点研发计划“七大农作物育种”重点专项(2016YFD01100201)

Research progress on genetic regulatory mechanism of seed color in soybean (Glycine max)

QIU Hong-Mei1(), CHEN Liang1, HOU Yun-Long1, WANG Xin-Feng1, CHEN Jian1, MA Xiao-Ping1, CUI Zheng-Guo1, ZHANG Ling1, HU Jin-Hai1, WANG Yue-Qiang1,*(), QIU Li-Juan2,*()   

  1. 1Jilin Academy of Agricultural Sciences / National Engineering Research Center for Soybean, Changchun 130033, Jilin, China
    2National Key Facility for Crop Gene Resources and Genetic Improvement (NFCRI) / Institute of Crop Sciences, Chinese Academy of Agricultural Sciences, Beijing 100081, China
  • Received:2021-01-24 Accepted:2021-05-17 Published:2021-12-12 Published online:2021-05-26
  • Contact: WANG Yue-Qiang,QIU Li-Juan
  • Supported by:
    National Key R&D Program for Crop Breeding(2016YFD01100201)

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摘要：

大豆种子颜色是重要的形态标记和进化性状, 在驯化过程中种皮从黑色逐渐演变成黄、绿、褐及双色, 子叶从绿色进化出黄色。深色种子中含有较多的天然色素——花色素, 具有药用和营养价值。因此, 种子颜色的遗传调控机制研究对进化理论和实际应用均具有重要意义。种子中色素含量及组分构成导致多样的种皮颜色, 其分子调控机制复杂。本文主要阐述了控制大豆种子颜色的遗传位点、相关基因与调控机制、类黄酮生物合成途径三方面的研究进展。具体介绍了9个经典遗传位点I、R、T、O、W1、K1、G、D1、D2和相关分子标记, 以及位点间的相互作用; 23个调控种子颜色的相关基因, 与部分基因等位变异的调控机制; 相关基因参与的类黄酮生物合成途径和主要代谢产物的生理功能。通过综述归纳了大豆种皮、种脐、子叶颜色的遗传调控研究进展, 利用遗传位点、基因、等位基因调控机制及类黄酮代谢途径绘制出调控网路, 以期为种子外观品质及花色苷组分遗传改良等研究提供参考。

关键词: 大豆, 种子颜色, 遗传调控, 相关基因, 类黄酮生物合成途径

Abstract:

The color of soybean seeds is an important morphological marker and evolutionary trait. During the process of domestication, seed coat has gradually evolved from black to yellow, green, black, brown, and bicolor, and cotyledons has evolved from green to yellow. The dark seed coat contains anthocyanins, which are natural pigments with medicinal and nutritional values. Therefore, it is of great importance to study the genetic regulation mechanism of seed color for evolutionary theory, variety breeding, and practical application. The pigment content and composition of seeds result in diverse seed coat colors through complex molecular regulatory mechanisms. In this paper, we described the research progress on genetic loci, related genes, regulatory mechanisms, and flavonoid biosynthesis pathways that controlling the color of soybean seeds. Specifically, we introduced the 9 classical genetic loci I, R, T, O, W1, K1, G, D1, D2, and related molecular markers, as well as the interactions between the loci; 22 related genes that controlling seed color, and the regulatory mechanisms of some allelic variants; as well as the physiological functions of the flavonoid biosynthesis pathways and major metabolites involved in the related genes. The progress of researches on genetic regulation of the color of seed coat, seed hilum, and cotyledon in soybean was reviewed, in addition the regulatory network was mapped with genetic loci, genes, allelic regulatory mechanisms, and flavonoid metabolic pathways, in order to provide references for the quality of seed appearance and genetic improvement of anthocyanin components.

Key words: soybean, seed color, genetic regulation, related gene, flavonoid biosynthesis pathway

表1

控制种子颜色经典遗传位点信息"

遗传位点
Genetic locus		 连锁群
Linkage group		 染色体
Chromosome		 控制性状
Character		 等位
基因
Allele		 表型
Phenotype		 参考文献
Reference
I A2 8 种皮、种脐是否含有花色素
The seed coat and hilum were controlled to contain anthocyanins.		 I 黄种皮黄脐
Fully yellow seed coat and hilum		 Terao[9], Todd et al.[26]
ii 黄种皮黑脐
Yellow seed coat with black hilum		 Terao[9], Tuteja et al.[27]
ik 鞍挂Black saddle Terao[9], Tuteja et al.[27]
i 黑种皮或褐种皮
Black or brown seed coat		 Terao[9], Clough et al.[28]
R K 9 种皮为黑色或褐色
The seed coat was black or brown.		 R 黑种皮Black seed coat Owen[10], Song et al.[29]
R* 黑种皮Black seed coat Zabala et al.[57]
r 褐种皮Brown seed coat Zabala et al.[57]
rm 虎斑Black spots or concentric rings of black pigment superimposed on an otherwise brown seed coat Zabala et al.[57]
T C2 6 种皮的着色程度及茸毛颜色
Control the pigmentation degree of seed coat and the color of trichomes.		 T 深褐种皮、棕色茸毛
Dark brown seed coat, tawny trichomes		 Owen[10], Zabala et al.[29,30]
t 褐种皮、灰色茸毛
Brown seed coat, gray trichomes		 Owen[10], Zabala et al.[29,30]
tm 深褐和褐色种皮、棕和灰色茸毛
Dark brown and brown seed coat, tawny and gray trichomes		 Zabala et al.[29,30]
O A2 8 种皮的褐色程度
Control the brown degree of seed coat.		 O 褐种皮Brown seed coat Owen[10], Yang et al.[32]
o 红褐种皮Red-brown seed coat Owen[10], Yang et al.[32]
W1 G 13 种皮着色程度及花的颜色
Control the pigmentation degree of seed coat and the color of flowers.		 W1 黑种皮、紫花
Black seed coat and purple flowers		 Owen[10], Zabala et al.[33]
w1 浅褐种皮、白花
Buff seed coat and white flowers		 Owen[10], Zabala et al.[33]
K1 B1 11 控制色素在鞍区的分布
Control the distribution of pigment in saddle region.		 K1 黄种皮Yellow seed coat Cho et al.[34]
k1 鞍挂Black saddle Cho et al.[34]
G D1a 1 种皮、子叶是否含有叶绿素
The seed coat and cotyledon were controlled to contain chlorophyll.		 G 绿种皮、绿子叶
Green seed coat and cotyledon		 Owen[10], Song et al.[21]
g 黄种皮、黄子叶
Yellow seed coat and cotyledon		 Owen[10], Song et al.[21]
D1 D1a 1 种皮、子叶是否含有叶绿素
The seed coat and cotyledon were controlled to contain chlorophyll.		 D1 黄种皮、黄子叶
Yellow seed coat and cotyledon		 Fang et al.[35], Song J [12]
d1 绿种皮、绿子叶
Green seed coat and cotyledon		 Fang et al.[35], Song J [12]
D2 B1 11 种皮、子叶是否含有叶绿素
The seed coat and cotyledon were controlled to contain chlorophyll.		 D2 黄种皮、黄子叶
Yellow seed coat and cotyledon		 Fang et al.[35], Song J [12]
d2 绿种皮、绿子叶
Green seed coat and cotyledon		 Fang et al.[35], Song J [12]
CytG — — 种皮、子叶是否含有叶绿素
The seed coat and cotyledon were controlled to contain chlorophyll.		 CytG 黄种皮、黄子叶
Yellow seed coat and cotyledon		 Guiamét et al.[39]
cytG 绿种皮、绿子叶
Green seed coat and cotyledon		 Guiamét et al.[39]

图1

不同基因型种子颜色 a: 黄种皮黄脐; b: 黄种皮黑脐; c: 鞍挂; d: 黑种皮; e: 褐种皮; f: 虎斑; g: 深褐种皮; h: 红褐种皮; i: 淡褐种皮; j: 绿种皮; k: 黄色子叶; l: 绿色子叶。"

图2

控制种子颜色遗传位点和相关分子标记 本图是在Song等[29]发表的遗传图谱基础上进行了修改。"

表2

调控种子颜色的相关基因信息"

序号
No.		 基因编号(v2.1)
Gene ID (v2.1)		 基因名称
Gene name		 英文缩写
Abbreviation		 遗传位点
Genetic locus		 编码产物注释
Annotation of coding products		 参考文献
Reference
1 Glyma.08G109500 查尔酮合成酶1
Chalcone synthase 1		 CHS1 I 类黄酮合成途径酶：合成柚皮苷查尔酮
Flavonoid biosynthesis pathway enzyme: synthesis of naringin chalcone		 Akada et al.[51]
2 Glyma.08G110300 查尔酮合成酶3
Chalcone synthase 3		 CHS3 I 类黄酮合成途径酶：合成柚皮苷查尔酮
Flavonoid biosynthesis pathway enzyme: synthesis of naringin chalcone		 Senda et al.[52]
Cho et al.[53]
3 Glyma.08G109200 查尔酮合成酶4
Chalcone synthase 4		 CHS4 I 类黄酮合成途径酶：合成柚皮苷查尔酮
Flavonoid biosynthesis pathway enzyme: synthesis of naringin chalcone		 Liu et al.[54]
Tuteja et al.[55]
4 Glyma.11G190900 Argonaute 5蛋白
Argonaute 5 protein		 AGO5 K1 调控CHS siRNAs在种皮的分布
Regulation of CHS siRNAs distribution in seed coat		 Rogers et al.[58]
Song et al.[59]
5 Glyma.09G025300 内啡肽酶同源物2a
Endorphinase homolog 2a		 DCL2a — 切割含长LIR的转录本,生成22nt siRNA
LIR-derived transcripts are predominately processed by GmDCL2 into 22-nt siRNAs		 Jia et al.[60]
6 Glyma.09G025400 内啡肽酶同源物2b
Endorphinase homolog 2a		 DCL2b — 切割含长LIR的转录本,生成22nt siRNA
LIR-derived transcripts are predominately processed by GmDCL2 into 22-nt siRNAs		 Jia et al.[60]
7 Glyma.06G202300 黄烷酮3’-羟化酶
Flavanone 3’-hydroxylase		 F3’H T 类黄酮合成途径酶：催化黄烷酮羟基化
Flavonoid biosynthesis pathway enzymes: catalyzing the hydroxylation of flavanones		 Toda et al.[61]
Guo et al.[62]
8 Glyma.13G072100 黄烷酮3’,5’-羟化酶
Flavanone 3’,5’-hydroxylase		 F3’5’H W1 类黄酮合成途径酶：催化黄烷酮羟基化
Flavonoid biosynthesis pathway enzymes: catalyzing the hydroxylation of flavanones		 Zabala et al.[33]
9 Glyma.08G062000 花青素还原酶
Anthocyanidin reductase		 ANR1 O 类黄酮合成途径酶：形成原花青素
Flavonoid biosynthesis pathway enzymes: procyanidins production		 Yang et al.[32]
10 Glyma.08G062100 花青素还原酶
Anthocyanidin reductase		 ANR2 O 类黄酮合成途径酶：形成原花青素
Flavonoid biosynthesis pathway enzymes: procyanidins production		 Yang et al.[32]
11 Glyma.09G235100 R2R3 MYB转录因子
Transcripion foctor R2R3 MYB		 R2R3 MYB R 转录因子：调控ANS基因的表达
Transcription factors: regulation of ANS gene expression		 Yan et al.[63]
Zabala et al.[64]
12 Glyma.01G198500 CAAX自身免疫蛋白酶
CAAX protease self-immunity		 CPSI G 防止叶绿素降解使成熟种皮保持绿色
Prevent chlorophyll degradation to keep the mature seed coat green		 Xie et al.[65]
Wang et al.[66]
13 Glyma.01g214600 保持绿色蛋白1
Protein stay-green 1		 SGR1 D1 使成熟种子保持绿色
Mature seeds stay-green		 Fang et al.[34]
宋健[12]
Fang et al.[34]
Song J[12]
14 Glyma.11g027400 保持绿色蛋白2
Protein stay-green 2		 SGR2 D2 使成熟种子保持绿色
Mature seeds stay-green		 Fang et al.[34]
宋健[12]
Fang et al.[34]
Song J[12]
15 Glyma.20G241500 查尔酮异构酶1A
Chalcone isomerase 1A		 CHI1A — 类黄酮合成途径酶：催化分子内环化反应
Flavonoid synthesis pathway enzymes: catalyzing intramolecular cyclization		 Kovinich et al.[63]
16 Glyma.20G241700 查尔酮异构酶2
Chalcone isomerase 2		 CHI2 — 类黄酮合成途径酶：催化分子内环化反应
Flavonoid synthesis pathway enzymes: catalyzing intramolecular cyclization		 Kovinich et al.[63]
17 Glyma.02G048400 黄烷酮3-羟化酶
Naringenin 3-dioxygenase		 F3H Wp 类黄酮合成途径酶：催化黄烷酮羟基化
Flavonoid biosynthesis pathway enzymes: catalyzing the hydroxylation of flavanones		 Zabala et al.[64]
18 Glyma.14G072700 二氢黄酮醇还原酶1
Dihydroflavonol 4-reductase 1		 DFR1 W3 类黄酮合成途径酶：形成无色花青素
Flavonoid synthesis pathway enzymes: formation of colorless anthocyanins		 Xu et al.[65]
19 Glyma.17G252200 二氢黄酮醇还原酶2
Dihydroflavonol 4-reductase 2		 DFR2 W4 类黄酮合成途径酶：形成无色花青素
Flavonoid synthesis pathway enzymes: formation of colorless anthocyanins		 Xu et al.[65]
20 Glyma.01G214200 花青素合成酶2
Anthocyanidin synthase 2		 ANS2 — 类黄酮合成途径酶：形成花色素
Flavonoid biosynthesis pathway enzymes: anthocyanin formation		 Kovinich et al.[68]
21 Glyma.11G027700 花青素合成酶3
Anthocyanidin synthase 3		 ANS3 — 类黄酮合成途径酶：形成花色素
Flavonoid biosynthesis pathway enzymes: anthocyanin formation		 Kovinich et al.[68]
22 Glyma.08G066800 UDP类黄酮葡萄糖基转移酶
UDP-flavonoid 3-O-glucosyltransferase		 UF3GT — 类黄酮合成途径酶：形成花色苷
Flavonoid biosynthesis pathway enzymes: anthocyanin formation		 Kovinich et al.[68]
23 — 光系统Ⅱ反应中心M亚基
Photosystem II reaction center M subunit		 Psbm — 具有叶绿素b降解功能
Degradation of chlorophyll b		 Kohzuma et al.[40]

表3

控制种子颜色等位基因的调控机制"

基因/编号
Gene ID		 突变基因型
Mutant genotype		 种子颜色
Seed color		 变异方式
Variation		 检测方法
Detection method		 参考文献
Reference
I
Glyma.08G109500
Glyma.08G110300
Glyma.08G109200		 ii 黄种皮黑脐 Yellow seed coat with black hilum CHS基因的拷贝数增加 The copy number of CHS gene increased BAC重测序 BAC resequencing Cho et al.[53]
ik 鞍挂 Black saddle 基因间序列缺失 Deletion of intergenic sequencesl BAC重测序 BAC resequencing Cho et al.[53]
i 深色种皮 Dark pigment seed coat 基因及基因间序列大片段缺失
Large deletions of genes and intergenic sequences		 BAC重测序 BAC resequencing Cho et al.[53]
R
Glyma.09G235100		 R* 黑种皮 Black seed coat 第2内含子插入CACTA转座子及CACTA序列的甲基化
Insertion of the second intron into the CACTA transposon and methylation of the CACTA sequence		 长片段PCR扩增、表达谱分析、甲基化检测
Long fragment PCR amplification, expression profile analysis, methylation detection		 Zabala et al.[64]
r 褐种皮 Brown seed coat 第2外显子单碱基C缺失 Single base C deletion in exon 2 克隆测序 Cloning and sequencing Zabala et al.[64]
rm 虎斑
Black spots or concentric rings of black pigment superimposed on an otherwise brown seed coat		 第2内含子插入CACTA转座子
Insertion of the second intron into the CACTA transposon		 长片段PCR扩增、表达谱分析、甲基化检测
Long fragment PCR amplification, expression profile analysis, methylation detection		 Zabala et al.[64]
T
Glyma.06G202300		 t 褐种皮 Brown seed coat 第3外显子单碱基A的缺失或非同义突变或单碱基C的缺失
Deletion of single base A or non synonymous mutation or deletion of single base C in exon 3		 PCR测序 PCR sequencing Guo et al.[62]
tm 深褐和褐色种皮
Dark brown and brown seed coat,		 第1内含子插入CACTA转座子
The first intron was inserted into the CACTA transposon		 长片段PCR扩增
Long fragment PCR amplification		 Zabala et al.[31]
W1
Glyma.13G072100		 w1 淡褐种皮 Buff seed coat 第3外显子插入53 bp
The third exon was inserted 53 bp sequence		 克隆测序
Cloning and sequencing		 Guo et al.[62]
K1
Glyma.11G190900		 k1 鞍挂 Black saddle 第7外显子129 bp的缺失
Deletion of 129 bp sequence in exon 7		 转录组与小RNA测序、PCR测序、全基因组深度重测序
Ranscriptome and micro RNA sequencing, PCR sequencing, whole genome deep resequencing		 Cho et al.[34]
G
Glyma.01G198500		 g 黄种皮 Yellow seed coat 第8内含子末端发生碱基突变
A base mutation occurred at the end of intron 8		 克隆测序
Cloning and sequencing		 Wang et al.[66]
D1
Glyma.01g214600		 d1 绿种皮、绿子叶
Green seed coat and cotyledon		 第2外显子单碱基T缺失
Single base T deletion in exon 2		 基因组重测序、克隆测序
Genome resequencing, cloning and sequencing		 Fang et al.[35], Song J[12]
D2
Glyma.11g027400		 d2 绿种皮、绿子叶
Green seed coat and cotyledon		 322 bp重复插入
322 bp sequence repeated insertion		 基因组重测序、克隆测序
Genome resequencing, cloning and sequencing		 Fang et al.[35], Song J[12]
CytG
cytG 绿种皮、绿子叶
Green seed coat and cotyledon		 5 bp插入 5 bp sequence insertion 对叶绿体基因组广泛的测序
Extensively sequenced the chloroplast genome		 Kohzuma et al.[40]

图3

种皮颜色相关基因调控的大豆类黄酮生物合成途径 修改自Zabala等[64]。"

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