浙江农业学报 ›› 2026, Vol. 38 ›› Issue (3): 609-620.DOI: 10.3969/j.issn.1004-1524.20250224
• 综述 • 上一篇
杜辰飞1,2(
), 李文珏1,2, 魏春艳2, 蔡丹英2, 王月志2, 施泽彬2, 戴美松2,*(
), 高永彬1,*(
)
收稿日期:2025-03-19
出版日期:2026-03-25
发布日期:2026-04-17
作者简介:戴美松,E-mail:daims@zaas.ac.cn通讯作者:
戴美松,高永彬
基金资助:
DU Chenfei1,2(
), LI Wenjue1,2, WEI Chunyan2, CAI Danying2, WANG Yuezhi2, SHI Zebin2, DAI Meisong2,*(
), GAO Yongbin1,*(
)
Received:2025-03-19
Published:2026-03-25
Online:2026-04-17
Contact:
DAI Meisong,GAO Yongbin
摘要:
在育种实践中,苹果、柑橘、梨等主要果树作物因其高度杂合的遗传背景、漫长的育种周期,以及数量性状的多基因调控特性,导致传统杂交育种效率受到限制。近年来,基于分子标记的高密度遗传图谱构建与数量性状基因座(QTL)精细定位技术已成为解析果树重要农艺性状遗传机制的关键手段。国内外已构建多张果树遗传连锁图谱,成功定位了与果实大小、形状、硬度、色泽、糖酸含量及香气等关键品质性状相关的QTL,并借助多组学整合分析揭示了其分子调控通路。本文系统综述了果树遗传图谱的发展现状和主要品质性状QTL定位的研究进展,重点指出:优化后的分子标记技术(SSR、SNP)显著提升了遗传图谱分辨率,“拟测交”与“双假测交”理论为QTL分析提供了关键方法框架;尽管目前已在多种果树中定位到主要品质性状相关QTL,但其稳定性与跨环境适应性仍需进一步验证;此外,多性状协同遗传机制,以及QTL与环境互作研究仍较薄弱,限制了标记辅助育种的精准性。未来应结合高通量测序、多组学与机器学习技术,构建超密遗传图谱并挖掘候选基因,为果树高效分子设计育种奠定理论基础。
中图分类号:
杜辰飞, 李文珏, 魏春艳, 蔡丹英, 王月志, 施泽彬, 戴美松, 高永彬. 果树遗传图谱构建与果实主要品质性状QTL定位研究进展[J]. 浙江农业学报, 2026, 38(3): 609-620.
DU Chenfei, LI Wenjue, WEI Chunyan, CAI Danying, WANG Yuezhi, SHI Zebin, DAI Meisong, GAO Yongbin. Research progress on genetic map construction in fruit trees and QTL identification of major fruit quality traits[J]. Acta Agriculturae Zhejiangensis, 2026, 38(3): 609-620.
| 物种 Species | 亲本 Parents | 标记类型 Marker types | 群体大小 (株) Population size (Number of plants) | 连锁群 数量 Number of linkage groups | 标记数量 Marker number | 平均标记 间距/cM Average marker spacing/cM | 构建年份 Construction year | 参考文献 References |
|---|---|---|---|---|---|---|---|---|
| 苹果 (Malus Mill.) | Gala×Jonathan | SNP/SSR | 145 | 17 | 765 | 1.96 | 2023 | [ |
| Fuji×Red3 | SNP | 168 | 17 | 630 | 0.30 | 2021 | [ | |
| Fuji×Hongrou | SSR/SRAP | 110 | 17 | 280 | 4.60 | 2016 | [ | |
| 柑橘 (Citrus L.) | MK×SB, Daisy | SNP | 165 | 9 | 2 588 | 0.54 | 2023 | [ |
| 梨橙2号×晚蜜2 | SSR/COS | 80 | 10 | 165 | 4.90 | 2017 | [ | |
| Licheng No. 2×Wanmi No. 2 | ||||||||
| Murcott tangor×Pera | DArT | 278 | 13 | 661 | 0.23 | 2017 | [ | |
| 桃 (Prunus L.) | 中油桃14号×黄水蜜 | In Del/SNP | 86 | 8 | 948 | 1.43 | 2019 | [ |
| Zhongyoutao No.14×Huangshuimi | ||||||||
| Zin Dai×Crimson Lady | SNP/SSR | 90 | 8 | 1 476 | 1.38 | 2018 | [ | |
| Venus×Big Top | SNP/SSR | 75 | 9 | 411 | 3.80 | 2016 | [ | |
| 梨 (Pyrus L.) | Niitaka×Hongxiangsu | SNP (Bin-Marker) | 176 | 17 | 3 190 | 0.43 | 2022 | [ |
| Red Clapp Favorite×Mansoo | SNP/SSR | 161 | 17 | 4 865 | 0.56 | 2017 | [ | |
| 八月红×砀山酥梨 | SNP/SSR | 102 | 17 | 3 241 | 0.70 | 2014 | [ | |
| Bayuehong×Dangshansu | ||||||||
| 葡萄 (Vitis L.) | B38×Horizon | rhAmpSeq | 118 | 19 | 1 092 | 1.01 | 2021 | [ |
| Horizon×Illinois 547-1 | rhAmpSeq | 142 | 19 | 1 171 | 1.16 | 2021 | [ | |
| Supreme×Nesbitt | GBS | 172 | 20 | 2 069 | 1.01 | 2019 | [ | |
| 枇杷 (Eriobotrya L.) | 宁海白×大丰 | SNP (Bin-Marker) | 130 | 17 | 3 859 | 0.52 | 2022 | [ |
| Ninghaibai×Dafeng | ||||||||
| Japonica×Deflexa | SNP | 96 | 17 | 960 | 1.78 | 2019 | [ | |
| 栎叶枇杷×解放钟 | RAD-seq/SRAP | 207 | 17 | 500 | 14.53 | 2017 | [ | |
| Quercifolia Loquat×Jiefangzhong | SLAF-seq/SSR |
表1 2014—2023年已发表的部分主要果树作物遗传连锁图谱
Table 1 Published genetic linkage maps of major fruit tree crops (2014—2023)
| 物种 Species | 亲本 Parents | 标记类型 Marker types | 群体大小 (株) Population size (Number of plants) | 连锁群 数量 Number of linkage groups | 标记数量 Marker number | 平均标记 间距/cM Average marker spacing/cM | 构建年份 Construction year | 参考文献 References |
|---|---|---|---|---|---|---|---|---|
| 苹果 (Malus Mill.) | Gala×Jonathan | SNP/SSR | 145 | 17 | 765 | 1.96 | 2023 | [ |
| Fuji×Red3 | SNP | 168 | 17 | 630 | 0.30 | 2021 | [ | |
| Fuji×Hongrou | SSR/SRAP | 110 | 17 | 280 | 4.60 | 2016 | [ | |
| 柑橘 (Citrus L.) | MK×SB, Daisy | SNP | 165 | 9 | 2 588 | 0.54 | 2023 | [ |
| 梨橙2号×晚蜜2 | SSR/COS | 80 | 10 | 165 | 4.90 | 2017 | [ | |
| Licheng No. 2×Wanmi No. 2 | ||||||||
| Murcott tangor×Pera | DArT | 278 | 13 | 661 | 0.23 | 2017 | [ | |
| 桃 (Prunus L.) | 中油桃14号×黄水蜜 | In Del/SNP | 86 | 8 | 948 | 1.43 | 2019 | [ |
| Zhongyoutao No.14×Huangshuimi | ||||||||
| Zin Dai×Crimson Lady | SNP/SSR | 90 | 8 | 1 476 | 1.38 | 2018 | [ | |
| Venus×Big Top | SNP/SSR | 75 | 9 | 411 | 3.80 | 2016 | [ | |
| 梨 (Pyrus L.) | Niitaka×Hongxiangsu | SNP (Bin-Marker) | 176 | 17 | 3 190 | 0.43 | 2022 | [ |
| Red Clapp Favorite×Mansoo | SNP/SSR | 161 | 17 | 4 865 | 0.56 | 2017 | [ | |
| 八月红×砀山酥梨 | SNP/SSR | 102 | 17 | 3 241 | 0.70 | 2014 | [ | |
| Bayuehong×Dangshansu | ||||||||
| 葡萄 (Vitis L.) | B38×Horizon | rhAmpSeq | 118 | 19 | 1 092 | 1.01 | 2021 | [ |
| Horizon×Illinois 547-1 | rhAmpSeq | 142 | 19 | 1 171 | 1.16 | 2021 | [ | |
| Supreme×Nesbitt | GBS | 172 | 20 | 2 069 | 1.01 | 2019 | [ | |
| 枇杷 (Eriobotrya L.) | 宁海白×大丰 | SNP (Bin-Marker) | 130 | 17 | 3 859 | 0.52 | 2022 | [ |
| Ninghaibai×Dafeng | ||||||||
| Japonica×Deflexa | SNP | 96 | 17 | 960 | 1.78 | 2019 | [ | |
| 栎叶枇杷×解放钟 | RAD-seq/SRAP | 207 | 17 | 500 | 14.53 | 2017 | [ | |
| Quercifolia Loquat×Jiefangzhong | SLAF-seq/SSR |
| 维度 Dimension | 果树 Fruit trees | 大田作物 Field crops |
|---|---|---|
| 群体类型 Group types | 多采用杂交F1群体或自然群体 Generally employ hybrid F1 populations or natural populations | 主要使用F2群体、RILs(重组自交系)或双单倍体群体 Mainly using F2 populations, RILs (recombinant inbred lines), or dihaploid populations |
| 群体规模 Group size | 规模较小(50~200株) Small scale (50-200 plants) | 规模较大(200~1 000株) Large scale (200-1 000 plants) |
| 标记类型 Marker type | 偏向SNP等高通量标记或SSR标记 Towards SNP and other high-throughput markers or SSR markers | SSR、AFLP等传统标记较为常用[ SSR, AFLP, and other traditional markers are more commonly used[ |
| 表型鉴定 Phenotypic identification | 需多年数据(3~5年) Requires multi-year data (3-5 years) | 单季/两年多点鉴定为主 Single season/over two years identification as main |
| 统计方法 Statistical methods | 需考虑多年数据混合模型,LOD阈值更高 Need to consider a multi-year data hybrid model with a higher LOD threshold | 复合区间作图法(CIM)为主,LOD值2.5~3.0[ Composite interval mapping (CIM) as the primary method, with LOD values ranging from 2.5 to 3.0[ |
| 环境影响 Environmental impact | 受环境影响大,环境互作效应显著 Highly influenced by the environment, with significant environmental interaction effects | 可通过多点重复降低误差[ Error can be reduced by multi-point repetition[ |
| 应用方向 Application direction | 侧重性状早期预测(成熟期、缩短童期)[ Focus on early prediction of traits (maturity period, shortened juvenile period)[ | 直接指导分子标记辅助选择[ Direct guidance in molecular marker-assisted selection[ |
表2 果树与大田作物QTL定位方法差异
Table 2 Differences in QTL mapping methods between fruit trees and field crops
| 维度 Dimension | 果树 Fruit trees | 大田作物 Field crops |
|---|---|---|
| 群体类型 Group types | 多采用杂交F1群体或自然群体 Generally employ hybrid F1 populations or natural populations | 主要使用F2群体、RILs(重组自交系)或双单倍体群体 Mainly using F2 populations, RILs (recombinant inbred lines), or dihaploid populations |
| 群体规模 Group size | 规模较小(50~200株) Small scale (50-200 plants) | 规模较大(200~1 000株) Large scale (200-1 000 plants) |
| 标记类型 Marker type | 偏向SNP等高通量标记或SSR标记 Towards SNP and other high-throughput markers or SSR markers | SSR、AFLP等传统标记较为常用[ SSR, AFLP, and other traditional markers are more commonly used[ |
| 表型鉴定 Phenotypic identification | 需多年数据(3~5年) Requires multi-year data (3-5 years) | 单季/两年多点鉴定为主 Single season/over two years identification as main |
| 统计方法 Statistical methods | 需考虑多年数据混合模型,LOD阈值更高 Need to consider a multi-year data hybrid model with a higher LOD threshold | 复合区间作图法(CIM)为主,LOD值2.5~3.0[ Composite interval mapping (CIM) as the primary method, with LOD values ranging from 2.5 to 3.0[ |
| 环境影响 Environmental impact | 受环境影响大,环境互作效应显著 Highly influenced by the environment, with significant environmental interaction effects | 可通过多点重复降低误差[ Error can be reduced by multi-point repetition[ |
| 应用方向 Application direction | 侧重性状早期预测(成熟期、缩短童期)[ Focus on early prediction of traits (maturity period, shortened juvenile period)[ | 直接指导分子标记辅助选择[ Direct guidance in molecular marker-assisted selection[ |
| QTL鉴定技术 QTL identification techniques | 优势 Advantages | 局限 Limitations | 参考文献 References |
|---|---|---|---|
| 遗传图谱QTL定位 Genetic map QTL mapping | 适用于主效QTL定位且结果稳定,辅助比较基因组分析 Applicable for major QTL localization with stable results, assisting in comparative genome analysis | 依赖高通量测序,成本较高;对群体类型要求严格,仅能解析亲本间有限的等位变异 Relies on high-throughput sequencing, which is costly; requires strict population types and can only resolve a limited number of allelic variations between parents | [ |
| RapMap高效克隆技术 RapMap high-efficiency cloning technology | 高效克隆主效QTL,适用于多亲本遗传多样性分析 Efficient cloning of major QTL for genetic diversity analysis in multi-parental populations | 依赖杂交群体构建,不适合自然群体,对微效QTL检测能力有限 Relies on hybrid population construction, not suitable for natural populations, and has limited capability in detecting minor QTL | [ |
| BSA(混池分离分析) BSA(bulked segregant analysis) | 成本低,无需完整遗传图谱,适合单一主效QTL分析 Low cost, without the need for a complete genetic map, suitable for the analysis of single major QTL | 分辨率较低,依赖极端表型材料;对候选区域后续验证工作量大 Low-resolution, relying on extreme phenotypic materials; substantial workload required for subsequent validation of candidate regions | [ |
| 多组学整合分析 Multi-omics integrated analysis | 适用于复杂性状(如风味、抗逆性)的多层次解析 Multilayered analysis for complex traits (such as flavor and stress resistance) | 数据量大、分析复杂、成本高,对样本处理和实验设计要求严格 Large data volume, complex analysis, high cost, and strict requirements for sample processing and experimental design | [ |
| GWAS(全基因组 关联分析) GWAS(genome-wide association study) | 可同时检测多个等位变异,适用于复杂性状 Can detect multiple allelic variants simultaneously, applicable to complex traits | 需大量样本控制假阳性;对微效QTL检测能力弱,且难以区分连锁位点 Requires a large number of samples to control false positives; weak detection capability for minor QTL and difficulty in distinguishing linked loci | [ |
表3 鉴定果树QTL方法及其优劣
Table 3 Methods for identifying QTL in fruit trees and their advantages and disadvantages
| QTL鉴定技术 QTL identification techniques | 优势 Advantages | 局限 Limitations | 参考文献 References |
|---|---|---|---|
| 遗传图谱QTL定位 Genetic map QTL mapping | 适用于主效QTL定位且结果稳定,辅助比较基因组分析 Applicable for major QTL localization with stable results, assisting in comparative genome analysis | 依赖高通量测序,成本较高;对群体类型要求严格,仅能解析亲本间有限的等位变异 Relies on high-throughput sequencing, which is costly; requires strict population types and can only resolve a limited number of allelic variations between parents | [ |
| RapMap高效克隆技术 RapMap high-efficiency cloning technology | 高效克隆主效QTL,适用于多亲本遗传多样性分析 Efficient cloning of major QTL for genetic diversity analysis in multi-parental populations | 依赖杂交群体构建,不适合自然群体,对微效QTL检测能力有限 Relies on hybrid population construction, not suitable for natural populations, and has limited capability in detecting minor QTL | [ |
| BSA(混池分离分析) BSA(bulked segregant analysis) | 成本低,无需完整遗传图谱,适合单一主效QTL分析 Low cost, without the need for a complete genetic map, suitable for the analysis of single major QTL | 分辨率较低,依赖极端表型材料;对候选区域后续验证工作量大 Low-resolution, relying on extreme phenotypic materials; substantial workload required for subsequent validation of candidate regions | [ |
| 多组学整合分析 Multi-omics integrated analysis | 适用于复杂性状(如风味、抗逆性)的多层次解析 Multilayered analysis for complex traits (such as flavor and stress resistance) | 数据量大、分析复杂、成本高,对样本处理和实验设计要求严格 Large data volume, complex analysis, high cost, and strict requirements for sample processing and experimental design | [ |
| GWAS(全基因组 关联分析) GWAS(genome-wide association study) | 可同时检测多个等位变异,适用于复杂性状 Can detect multiple allelic variants simultaneously, applicable to complex traits | 需大量样本控制假阳性;对微效QTL检测能力弱,且难以区分连锁位点 Requires a large number of samples to control false positives; weak detection capability for minor QTL and difficulty in distinguishing linked loci | [ |
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