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作物学报 ›› 2026, Vol. 52 ›› Issue (6): 1711-1727.doi: 10.3724/SP.J.1006.2026.51096

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

小麦ABHD6基因家族鉴定与粒重功能标记开发

张献丰1,2(), 郭利建1, 李康春1,2, 孔斌雪3, 刘玉芳1,2, 车卓1, 杨德龙1,2,3,*()   

  1. 1 甘肃农业大学干旱生境作物学国家重点实验室, 甘肃兰州 730070
    2 甘肃农业大学生命科学技术学院, 甘肃兰州 730070
    3 甘肃农业大学农学院, 甘肃兰州 730070
  • 收稿日期:2025-11-27 接受日期:2026-03-16 出版日期:2026-06-12 网络出版日期:2026-03-30
  • 通讯作者: * 杨德龙, E-mail: yangd1@gsau.edu.cn
  • 作者简介:张献丰, E-mail: 3029202284@qq.com
  • 基金资助:
    甘肃省科技重大专项(25ZDNA001);甘肃省麦类产业技术体系项目(GSARS07);甘肃省教育厅青年博士支持项目(2024QB-062);省部共建干旱生境作物学国家重点实验室开放基金项目(GSCS-2023-07)

Identification of the ABHD6 gene family and development of functional markers for grain weight in wheat

Zhang Xian-Feng1,2(), Guo Li-Jian1, Li Kang-Chun1,2, Kong Bin-Xue3, Liu Yu-Fang1,2, Che Zhuo1, Yang De-Long1,2,3,*()   

  1. 1 State Key Laboratory of Aridland Crop Science, Gansu Agricultural University, Lanzhou 730070, Gansu, China
    2 College of Life Science and Technology, Gansu Agricultural University, Lanzhou 730070, Gansu, China
    3 College of Agronomy, Gansu Agricultural University, Lanzhou 730070, Gansu, China
  • Received:2025-11-27 Accepted:2026-03-16 Published:2026-06-12 Published online:2026-03-30
  • Contact: * Yang De-Long, E-mail: yangd1@gsau.edu.cn
  • Supported by:
    Gansu Provincial Science and Technology Major Project(25ZDNA001);Gansu Province Wheat Industry Technology System Project(GSARS07);Youth Doctoral Support Project of Gansu Provincial Department of Education(2024QB-062);Open Fund of the State Key Laboratory of Cropology in Arid Habitat Jointly Established by the Province and Ministry(GSCS-2023-07)

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摘要：

α/β-水解酶蛋白6 (α/β-hydrolasedomain-containing 6, ABHD6)作为丝氨酸水解酶超家族的重要成员, 在调控植物生长发育及环境胁迫响应中发挥重要作用。然而, 目前仍缺乏对小麦(Triticum aestivum L.) ABHD6基因家族的系统性鉴定与分析。本研究综合利用系统进化、蛋白质序列分析、启动子顺式作用元件预测以及单倍型分析等方法, 在小麦基因组中共鉴定出18个TaABHD6家族成员。系统进化与结构分析表明, 小麦TaABHD6蛋白与水稻OsABHD6蛋白亲缘关系较近, 同一进化分支内的成员具有高度相似的保守结构域。启动子分析显示, TaABHD6基因家族启动子区富含多种参与植物生长发育调控的顺式作用元件。通过分析TaABHD6基因家族成员的基因组多态性及其与籽粒表型的关联, 鉴定出8个基因存在不同的单倍型。其中, 3个基因的不同单倍型与籽粒相关性状之间存在显著关联。TaABHD6-5基因的不同单倍型与粒重、粒长、粒宽和粒厚均显著关联。针对TaABHD6-5启动子区域-1045 bp处SNP (A/T)开发KASP分子标记, 在305份小麦种质中进行验证, 发现该功能标记可有效区分2种单倍型, 其中, TaABHD6-5-HapI单倍型品种268份, TaABHD6-5-HapII单倍型品种34份, 杂合子2份, 分型失败1份。籽粒表型数据关联分析表明, TaABHD6-5-HapI是增加粒重的优异单倍型。表达模式分析结果表明, TaABHD6-5在孕穗期的茎、穗以及籽粒发育初期表达较高, 且优异单倍型TaABHD6-5-HapI的表达量显著低于TaABHD6-5-HapII, 表明TaABHD6-5可能为粒重负调控因子。基因序列分析表明, TaABHD6-5受miR160靶向调控。本研究结果为阐明TaABHD6基因调控小麦籽粒发育的生物学功能及其优异单倍型的应用价值提供重要理论支撑。

关键词: 小麦, α/β-水解酶结构域蛋白6 (ABHD6), 基因家族鉴定, 单倍型, 分子标记

Abstract:

α/β-Hydrolase domain-containing protein 6 (ABHD6), a key member of the serine hydrolase superfamily, plays important roles in regulating plant growth, development, and responses to environmental stress. However, a systematic genome-wide identification and analysis of the ABHD6 gene family in wheat (Triticum aestivum L.) is still lacking. Here, we integrated phylogenetic analysis, protein sequence characterization, promoter cis-acting element prediction, and haplotype analysis, and identified 18 TaABHD6 family members in the wheat genome. Phylogenetic and structural analyses showed that TaABHD6 proteins are closely related to rice OsABHD6 proteins, and that members within the same clade share highly conserved domains. Promoter analysis revealed that TaABHD6 promoters are enriched in cis-acting elements associated with plant growth and development. By analyzing polymorphisms in TaABHD6 genes and their associations with grain phenotypes, we identified eight genes with distinct haplotypes. Haplotype variation in three genes was significantly associated with grain-related traits; in particular, TaABHD6-5 haplotypes were significantly associated with grain weight, length, width, and thickness. We developed a KASP marker based on an A/T SNP at -1045 bp in the TaABHD6-5 promoter. Validation in 305 wheat accessions showed that this functional marker effectively distinguished two haplotypes: 268 accessions carried TaABHD6-5-HapI, 34 carried TaABHD6-5-HapII, two were heterozygous, and one accession failed genotyping. Association analysis further indicated that TaABHD6-5-HapI is a favorable haplotype for increased grain weight. qRT-PCR showed that TaABHD6-5 was relatively highly expressed in stems and spikes at booting and during early grain development; notably, expression was significantly lower in the favorable haplotype TaABHD6-5-HapI than in TaABHD6-5-HapII, suggesting that TaABHD6-5 may act as a negative regulator of grain weight. Sequence analysis further suggested that TaABHD6-5 is targeted by miR160. Together, these results provide genome-wide evidence for the roles of TaABHD6 genes in wheat grain development and highlight the potential breeding value of superior haplotypes.

Key words: wheat, α/β-hydrolase domain-containing protein 6 (ABHD6), gene family identification, haplotype, molecular marker

表1

序列号对应品种名称"

序列号
Serial number		 单倍型信息
Haploid information		 品种名称
Variety name
XN30 HapI 11-257-4-1
XN37 HapI 1389-2
XN38 HapI 1-4-8-1
XN41 HapI 1R14
XN46 HapII 1R25
XN68 HapI 99384-2-1
XN151 HapII 复壮30 Fuzhuang 30
XN156 HapII 旱选10号 Hanxuan 10
XN171 HapII 衡95观26 Heng 95 Guan 26
XN203 HapII 冀麦22 Jimai 22

表2

本研究所用引物"

引物名称
Primer name		 引物序列
Primer sequence (5′-3′)
TaABHD6-5 F: CGAAGCTCATAGGCGCGA
R: GATCACCACCCGCTCCAC
TaACTIN F: AGGTGCCCTGAGGTGCTGTT
R: GCCAAAATAGAGCCACCGAT

表3

单倍型表达模式分析所用引物"

引物名称
Primer name		 引物序列
Primer sequence (5′-3′)
KASP-TaABHD6-F1 GAAGGTCGGAGTCAACGGATTCGTGCCTGTTTGCGAGTATA
KASP-TaABHD6-F2 GAAGGTGACCAAGTTCATGCTCGTGCCTGTTTGCGAGTATT
KASP-TaABHD6-R CTTGACAGTTGATGAATAGATCGG

表4

小麦TaABHD6基因家族"

基因名称
Gene name		 基因号
Gene ID		 氨基酸数量
No. of amino acids (aa)		 相对分子量Molecular weight (kD) 等电点
Isoelectric point		 不稳定指数Instability
index		 亲水性
Hydrophilicity		 亚细胞定位
Subcellular localization
TaABHD6-1 TraesCS1A02G435000.1 308 32.86 4.56 36.73 0.118 cyto
TaABHD6-2 TraesCS1B02G470800.1 309 32.87 4.64 38.74 0.175 cyto
TaABHD6-3 TraesCS1D02G444100.1 334 35.89 4.54 38.13 0.105 cyto
TaABHD6-4 TraesCS2A02G163300.1 332 36.94 8.80 48.61 0.023 cyto
TaABHD6-5 TraesCS2A02G171000.1 275 30.20 8.81 51.31 -0.036 chlo
TaABHD6-6 TraesCS2B02G189100.1 334 37.20 7.72 48.17 0.025 cyto
TaABHD6-7 TraesCS2B02G197400.1 314 34.67 7.82 53.02 -0.050 chlo
TaABHD6-8 TraesCS2D02G170200.1 334 37.09 8.58 49.18 -0.013 cyto
TaABHD6-9 TraesCS2D02G178700.1 314 34.59 7.03 49.82 -0.051 chlo
TaABHD6-10 TraesCS3A02G538800.2 260 28.92 5.19 33.54 -0.136 cyto
TaABHD6-11 TraesCS3D02G545700.2 260 29.10 5.29 28.91 -0.180 cyto
TaABHD6-12 TraesCS4A02G010600.1 324 35.56 5.92 45.22 -0.043 chlo
TaABHD6-13 TraesCS4B02G293800.1 307 33.62 6.08 46.19 -0.100 chlo
TaABHD6-14 TraesCS4B02G293900.1 325 35.24 7.78 41.47 -0.030 chlo
TaABHD6-15 TraesCS4D02G292500.1 322 35.50 6.09 49.03 -0.108 chlo
TaABHD6-16 TraesCS4D02G292600.1 322 35.29 5.54 42.11 -0.028 chlo
TaABHD6-17 TraesCS4D02G292700.1 320 34.92 8.54 38.01 -0.031 cyto
TaABHD6-18 TraesCS5D02G425500.1 227 25.53 9.01 47.97 -0.025 extr

图1

小麦TaABHD6基因家族在染色体上的定位"

图2

ABHD6基因家族系统进化树分析 不同颜色代表ABHD6基因家族的不同亚族。淡紫色圆形表示bootstrap值; 矩形代表小麦; 深紫色圆形代表大麦; 五角星代表玉米; 红色三角形代表水稻; 蓝色三角形代表拟南芥。"

图3

小麦 TaABHD6基因家族成员的进化关系、保守基序和基因结构"

图4

小麦TaABHD6基因家族共线性分析 A: TaABHD6基因家族种内共线性分析; B: TaABHD6基因家族种间共线性分析; 染色体标号前缀Ta代表普通小麦, Os代表水稻。"

表5

小麦TaABHD6基因家族Ka/Ks分析"

基因1
Gene 1		 基因2
Gene 2		 非同义替换率
Ka		 同义替换率
Ks		 非同义替换率/同义替换率
Ka/Ks
TaABHD6-1 TaABHD6-2 0.029560476405657555 0.14576965151507500 0.20278896257497406
TaABHD6-1 TaABHD6-3 0.040760832579004340 0.13490226857082155 0.30215083119677527
TaABHD6-2 TaABHD6-3 0.030917036867965757 0.11045957175281135 0.27989459290275470
TaABHD6-4 TaABHD6-6 0.030353271379501870 0.06159789458113049 0.49276475415119300
TaABHD6-5 TaABHD6-7 0.014547673945103423 0.08534953822776796 0.17044818574508028
TaABHD6-4 TaABHD6-8 0.020418683960993462 0.07088605521155301 0.28804937586180734
TaABHD6-5 TaABHD6-9 0.021896701808340460 0.10876786590712034 0.20131590912189648
TaABHD6-5 TaABHD6-15 0.381523080690999900 1.00018318421061040 0.38145320448685116
TaABHD6-6 TaABHD6-8 0.028574884674202803 0.08796681489232491 0.32483709577503360
TaABHD6-7 TaABHD6-9 0.013050111753615083 0.07239658798238977 0.18025865744929140
TaABHD6-7 TaABHD6-15 0.382730436542767500 0.93268675733376540 0.41035260073474594

图5

小麦TaABHD6基因家族启动子顺式作用元件"

图6

小麦TaABHD6基因家族结构及序列多态性分析和不同单倍型的关联分析 A-C分别表示小麦基因TaABHD6-1、TaABHD6-5和TaABHD6-7结构及序列多态性分析; D-F分别表示小麦基因TaABHD6-1、TaABHD6-5和TaABHD6-7不同单倍型的关联分析。横坐标表示田间试验的3个年份(2002、2005和2010年)。"

图7

TaABHD6基因家族单倍型与籽粒性状的关联分析 A: TaABHD6-5的KASP基因分型散点聚类图; B: 籽粒相关性状与TaABHD6-5不同单倍型的关联分析。红点、绿点和蓝点分别表示TaABHD6-5-HapI纯合子TT、杂合子AT、TaABHD6-5-HapII纯合子AA; E1-E5分别代表2020年通渭、2021年通渭旱地、2021年通渭水地、2022年通渭和2022年庄浪。"

图8

小麦TaABHD6-5时空表达模式及单倍型表达差异分析 A: TaABHD6-5在小麦不同组织及不同籽粒发育时期的表达模式; 5、10、15 和20 d分别代表授粉后第5、10、15和20天的籽粒; B: TaABHD6-5在2种不同单倍型间的表达水平差异分析; **表示与TaABHD6-5-HapI相比差异极显著(P < 0.01); 用于不同单倍型表达量分析的材料包括5份TaABHD6-5-HapI品种(XN30、XN37、XN38、XN41、XN68)和5份TaABHD6-5-HapII品种(XN46、XN151、XN156、XN171、XN203)。"

图9

TaABHD6-5序列调控元件分析 A: 小麦TaABHD6-5上游miRNA的生物信息学预测; B: tae-miR160介导的TaABHD6-5转录本切割的降解组分析。"

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