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作物学报 ›› 2026, Vol. 52 ›› Issue (3): 708-721.doi: 10.3724/SP.J.1006.2026.55055

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

甘蓝型春油菜早花基因BnCRY2功能位点KASP标记的开发及应用

王楚锐(), 李开祥, 赵志, 肖麓, 唐国永, 赵志刚, 徐亮, 杜德志, 柳海东()   

  1. 青海大学农林科学院 / 国家油菜改良中心青海分中心 / 青海省春油菜工程技术研究中心 / 青海省春油菜遗传改良重点实验室 / 农业农村部春油菜科学观测实验站, 青海西宁 810016
  • 收稿日期:2025-08-11 接受日期:2025-11-18 出版日期:2026-03-12 网络出版日期:2025-11-28
  • 通讯作者: *柳海东, E-mail: dahaima@163.com
  • 作者简介:E-mail: waqhy1121@163.com
  • 基金资助:
    国家自然科学基金项目(32260507);农业生物育种重大专项(2022ZD04009);财政部和农业农村部国家现代农业产业技术体系建设专项(CRAS-12);青海省帅才科学家负责制项目(2022-NK-170)

Development and application of KASP markers for functional sites of the early flowering gene BnCRY2 in Brassica napus L. spring rapeseed

Wang Chu-Rui(), Li Kai-Xiang, Zhao Zhi, Xiao Lu, Tang Guo-Yong, Zhao Zhi-Gang, Xu Liang, Du De-Zhi, Liu Hai-Dong()   

  1. Academy of Agricultural and Forestry Sciences, Qinghai University / Qinghai Research Branch of the National Rapeseed Genetic Improvement Center / Qinghai Spring Rape Engineering Research Center / Key Laboratory of Qinghai Province for Spring Rapeseed Genetic Improvement / Spring Rape Scientific Observation Experimental Station of Ministry of Agriculture and Rural Areas, Xining 810016, Qinghai, China
  • Received:2025-08-11 Accepted:2025-11-18 Published:2026-03-12 Published online:2025-11-28
  • Contact: *柳海东, E-mail: dahaima@163.com
  • Supported by:
    National Natural Science Foundation of China(32260507);Major Project of Agricultural Biological Breeding(2022ZD04009);China Agriculture Research System of MOF and MARA(CRAS-12);Qinghai Province Leading Scientist Responsibility System Project(2022-NK-170)

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摘要：

通过前期对早花位点BnaC08cqDTF的候选基因BnCRY2的克隆发现, 其等位基因第3外显子存在一个A/G的SNP, 根据该SNP开发了相应的KASP标记BN900449, 进一步利用上述标记及与早花位点qFT.A02-1和qFTA10紧密连锁的4个标记(多重PCR标记PB05、PB06和KASP标记A015348、K000272)对497份甘蓝型油菜自然群体进行早花基因型鉴定, 结果显示, 3个位点均可将自然群体分成花期显著差异的2组, 且含双位点品系的平均开花时间早于单位点品系; 从中筛选出分别含有BnaC08cqDTF、qFTA10、qFT.A02-1位点的资源各3份, 不同单位点资源两两杂交进行位点聚合, 结合分子标记辅助选择创建特早熟资源41份, 这些资源比亲本早开花3~4 d; 从中筛选出初花期较早且品质双低的品系195, 连续2年在高海拔区5点进行测产试验, 结果表明, 2年平均产量比高海拔春油菜区白菜型主栽品种浩油11号增产23.07%。该研究不仅拓宽了特早熟甘蓝型春油菜种质资源创建的思路, 也为特早熟高产优质油菜品种的培育提供了亲本支撑。

关键词: 基因聚合, 甘蓝型春油菜, 早花资源, 分子标记, 早花位点

Abstract:

Cloning of the candidate gene BnCRY2, associated with the early-flowering locus BnaC08cqDTF, identified an A/G single-nucleotide polymorphism (SNP) within the third exon of the corresponding alleles. Based on this SNP, a KASP marker named BN900449 was developed. This marker, along with four additional markers—multiplex PCR markers PB05 and PB06, and KASP markers A015348 and K000272—tightly linked to the early-flowering loci qFT.A02-1 and qFTA10, was used to genotype a natural population of 497 Brassica napus L. accessions. Genotypic analysis revealed that each of the three loci effectively divided the population into two groups with significantly different flowering times. Notably, accessions carrying two of the early-flowering loci flowered, on average, earlier than those carrying only one. Three representative accessions, each harboring a different single-locus allele (BnaC08cqDTF, qFTA10, and qFT.A02-1), were selected for diallel crosses aimed at pyramiding early-flowering alleles. Marker-assisted selection subsequently yielded 41 extremely early-flowering lines, which exhibited flowering times 3-4 days earlier than their respective parents. Among these, 195 lines were identified that combined extreme earliness with double-low seed quality. Multi-location yield trials conducted at five high-altitude sites over two consecutive years showed that the average yield of selected lines exceeded that of Brassica rapa L. cultivar Haoyou 11—the predominant spring-sown variety in high-altitude regions—by 23.07%. This study not only expands the genetic framework for developing extremely early- flowering B. napus spring-type germplasm but also provides elite parental resources for breeding early-maturing, high-yielding, and high-quality rapeseed cultivars.

Key words: gene pyramiding, spring Brassica napus L., early flowering resources, molecular marker, early flowering locus

表1

KASP及靶向测序引物序列信息"

位点信息
Locus
information		 引物Primer 位置
Position
(bp)		 突变Mutation 上游引物序列
Forward primer
sequence
(5°-3°)		 下游引物序列
Reverse primer
sequence
(5°-3°)		 通用引物序列
Universal primer sequence
(5°-3°)		 来源
Source
qFTA10 A015348 20,934,635 A/T GAAGGTGACCAAGTTCATGCGGCTCAGTCGATTCATAATGCATC GAAGGTCGGAGTCAACGGATTGCGCGGCTCAGTCGATTCATAATGCATA CTGAAGGAGCTATTCTGATCCCCAA 数据未发表
The data are unpublished
K000272 20,874,901 A/T GAAGGTGACCAAGTTCATGCTAGTGTTATGATTCCATTGCGCAAACA GAAGGTCGGAGTCAACGGATTAGTGTTATGATTCCATTGCGCAAACT CTCTGCAAGTCCTTGTTTTCATCAAGATT
qFT.A02-1 PB05 7,420,167 A/T CCTACACGACGCTCTTCCGATCTTTTAGTAGAAGAAGAAGAGGTGGATGATCTTG TCAGACGTGTGCTCTTCCGATCTAAACCCTTTGTACATCTCTCCCCAATCTAT Li, et al[4]
PB06 7,444,182 G/A CCTACACGACGCTCTTCCGATCTATAATACATAATTCGAGAAGCAAACCCATCAA TCAGACGTGTGCTCTTCCGATCTATTTTGACAGTATTTTGAACCGCTTTTTATCC

表2

自然群体初花期表型统计分析表"

材料名称
Material name		 资源数
Number of resources		 初花期First flowering period (d)
95%置信区间下限
Lower limit of the 95% confidence interval		 95%置信区间上限
Upper limit of the 95% confidence interval		 变幅
Range		 平均值±标准差
Mean ± SD
自然群体
Natural population		 497 49.58 50.76 33.67-74.50 50.17±6.66

表3

早花基因BnCRY2 A/G突变的KASP标记序引物序列信息"

位点候选
基因信息
Locus candidate gene information		 引物Primer 位置
Position (bp)		 突变、插入
/缺失Mutate, insert / miss		 上游引物序列
Forward primer sequence
(5′-3′)		 下游引物序列
Reverse primer sequence
(5′-3′)		 通用引物序列
Universal primer sequence (5′-3′)
BnaC08cqDTF BN900449 903326 C/T* GAAGGTGACCAAGTTCATGCTCCCAAAAATATTCATCCCCCACTC GAAGGTCGGAGTCAACGGATTCCCAAAAATATTCATCCCCCACTT TACTGGATGGATGCATAACAGGATT

表4

497份甘蓝型春油菜资源在三早花位点5个标记中基因型鉴定结果"

基因型Genotype 资源数
Number of resources		 初花期First flowering period (d) P-value
qFT.A02-1 qFTA10 BnaC08cqDTF 变幅
Range		 平均值±标准差
Mean ± SD
PB05 PB06 K000272 A015348 BN900449
AA AA 224 33.67-68.67 49.03±7.04 2.0×10-5
BB BB 223 37.33-74.50 51.64±5.67
AA AA 209 34.00-65.67 48.89±6.63 3.4×10-5
BB BB 203 33.67-74.50 51.55±6.22
AA
BB		 321 34.00-74.50 49.11±6.45 5.0×10-6
155 33.67-71.50 52.08±6.80

图1

5个标记基因分型图 图A为标记BN900449基因分型图; 图B为标记A015348基因分型图; 图C为标记K000272基因分型图; 图D为标记PB05基因分型图;图E为标记PB06基因分型图。图中蓝色部分为含有早花位点纯合的基因型, 标记红色部分为不含早花位点纯合基因型, 紫色部分为含有早花位点杂合基因型。FAM为与其中一个等位基因特异性结合的荧光染料, VIC为与另一个等位基因特异性结合的荧光染料。"

图2

497份甘蓝型春油菜资源在三早花位点5个标记中花期分布及显著关系 图中AA基因型均代表含有该位点早花基因的基因型, BB基因型均代表不含该位点早花基因的基因型, *表示经过t检验, 所指基因型之间存在显著差异(P < 0.05), **表示经过t检验, 所指基因型之间存在极显著差异(P < 0.01)。"

表5

497份甘蓝型春油菜资源在三早花位点5个标记中含有双早花位点的基因型鉴定结果"

基因型Genotype 资源数
Number of resources		 初花期First flowering period (d) P-value
qFT.A02-1 qFTA10 BnaC08cqDTF 变幅
Range		 平均值±标准差
Mean ± SD
PB05 PB06 K000272 A015348 BN900449
AA AA AA AA 94 34.00-64.00 47.19±7.30 2.0×10-6
BB BB BB BB 97 37.33-74.50 52.03±6.41
AA AA AA 132 34.00-64.00 47.28±6.30 3.0×10-6
BB BB BB 55 33.67-71.50 52.28±6.74
AA AA AA 127 34.00-65.50 46.79±6.52 2.9×10-9
BB BB BB 50 37.33-71.50 53.05±6.30

图3

5个标记在亲本及F2、F5中的基因分型图 图A, B, C为亲本在5个标记下鉴定结果, 其中A为BnaC08cqDTF基因功能标记的鉴定结果, B为qFTA10两个标记的鉴定结果, C为qFT.A02-1两个标记的鉴定结果。图D, E, F为F2代在5个标记下的鉴定结果, 其中D为qFT.A02-1两个标记的鉴定结果, E为qFTA10两个标记的鉴定结果, F为BnaC08cqDTF基因功能标记的鉴定结果。图G, H, I为F5代在5个标记下的鉴定结果, 其中G为qFT.A02-1两个标记的鉴定结果, H为qFTA10两个标记的鉴定结果, I为BnaC08cqDTF基因功能标记的鉴定结果。图A, F, I针对BnaC08cqDTF基因功能标记, C:C基因型是含有早花位点的纯合基因型, C:T基因型是含有早花位点的杂合基因型, T:T基因型是不含早花位点的纯合基因型, 图B, C, D, E, G, H针对不同标记AA基因型皆代表含有早花位点纯合基因型, AB基因型皆代表含有早花位点杂合基因型, BB基因型皆代表不含早花位点纯合基因型。"

表6

单位点亲本及位点聚合系开花时间表现与品质"

材料名称
Material name		 位点
Locus		 来源
Source		 开花时间
Flowering time
(d)		 芥酸
Mustard (%)		 硫苷
Thioside
(μmol g-1)		 含油量
Oil content (%)
N80 qFTA10 43.20±0.84 a 0.29 0 39.82
N78 qFTA10 37.20±1.30 ef 0.06 2.21 33.57
N108 qFTA10 37.60±1.14 de 1.23 29.00 38.07
N64 qFT.A02-1 36.20±1.79 fg 0 75.21 38.73
N67 qFT.A02-1 37.80±0.84 de 0.24 0 30.99
N103 qFT.A02-1 35.40±1.34 gh 1.48 11.33 42.89
N62 BnaC08cqDTF 43.80±0.84 a 5.32 5.89 39.95
N74 BnaC08cqDTF 41.80±1.30 b 0 0 36.94
N133 BnaC08cqDTF 41.40±1.14 b 1.03 10.37 41.14
12 qFT.A02-1&qFTA10 N103 × N78 32.60±0.55 jk 0 5.30 41.08
51 qFT.A02-1&qFTA10 N64 × N80 33.40±0.55 jk 0 5.41 38.43
60 qFT.A02-1&qFTA10 N64 × N78 33.40±0.55 jk 0.51 11.69 40.39
63 qFT.A02-1&qFTA10 N64 × N78 33.40±0.55 jk 0.30 16.83 35.97
293 qFT.A02-1&qFTA10 N67 × N80 34.20±0.45 hij 0.12 1.26 42.19
328 qFT.A02-1&qFTA10 N103 × N108 34.00±0.71 hijk 0 13.33 40.45
329 qFT.A02-1&qFTA10 N103 × N108 33.80±0.45 ijk 3.14 178.43 27.02
330 qFT.A02-1&qFTA10 N103 × N108 33.40±0.55 jk 1.45 136.31 27.62
335 qFT.A02-1&qFTA10 N103 × N108 33.40±0.55 jk 2.76 155.11 34.11
65 qFT.A02-1&BnaC08cqDTF N103 × N133 33.40±0.55 jk 2.79 144.25 27.61
67 qFT.A02-1&BnaC08cqDTF N103 × N133 34.00±1.00 hijk 0.99 50.08 37.42
76 qFT.A02-1&BnaC08cqDTF N103 × N133 33.80±0.84 ijk 1.95 58.37 38.58
243 qFT.A02-1&BnaC08cqDTF N67 × N62 34.20±0.45 hij 0.69 44.02 40.67
371 qFT.A02-1&BnaC08cqDTF N67 × N74 34.00±0.71 hijk 3.97 176.86 34.03
374 qFT.A02-1&BnaC08cqDTF N67 × N74 33.80±0.45 ijk 4.06 220.07 36.81
375 qFT.A02-1&BnaC08cqDTF N67 × N74 34.40±1.82 hij 4.29 191.62 35.66
425 qFT.A02-1&BnaC08cqDTF N67 × N62 36.20±1.30 fg 0.99 123.04 37.58
430 qFT.A02-1&BnaC08cqDTF N67 × N62 34.40±1.67 hij 2.28 80.35 30.09
138 qFTA10&BnaC08cqDTF N78 × N74 34.20±0.45 hij 0.09 14.80 39.88
140 qFTA10&BnaC08cqDTF N78 × N74 34.20±0.45 hij 1.42 70.01 42.54
142 qFTA10&BnaC08cqDTF N78 × N74 33.60±0.55 ijk 0 7.26 35.30
146 qFTA10&BnaC08cqDTF N78 × N74 34.40±0.55 hij 0 11.96 41.93
148 qFTA10&BnaC08cqDTF N78 × N74 34.20±0.84 hij 2.50 126.25 23.23
151 qFTA10&BnaC08cqDTF N78 × N74 34.40±0.55 hij 0 2.04 38.80
152 qFTA10&BnaC08cqDTF N78 × N74 34.20±0.45 hij 0.57 39.16 29.40
153 qFTA10&BnaC08cqDTF N78 × N74 34.40±0.55 hij 0 3.02 31.16
158 qFTA10&BnaC08cqDTF N78 × N74 33.60±0.55 ijk 0 21.57 31.33
165 qFTA10&BnaC08cqDTF N108 × N62 33.60±0.55 ijk 1.94 79.56 40.05
181 qFTA10&BnaC08cqDTF N80 × N62 38.40±0.89 cde 0 7.46 40.60
182 qFTA10&BnaC08cqDTF N80 × N62 39.20±0.84 c 4.19 155.46 27.25
183 qFTA10&BnaC08cqDTF N80 × N62 38.40±0.55 cde 0.04 18.89 43.17
209 qFTA10&BnaC08cqDTF N80 × N74 34.20±0.45 hij 3.21 56.23 44.63
200 qFTA10&BnaC08cqDTF N80 × N74 34.60±1.34 hij 0.02 10.67 41.28
205 qFTA10&BnaC08cqDTF N80 × N74 34.40±0.89 hij 0.76 10.83 41.43
195 qFTA10&BnaC08cqDTF N78 × N133 33.80±0.45 ijk 0.03 18.37 46.28
217 qFTA10&BnaC08cqDTF N78 × N133 38.60±0.89 cd 0 8.91 41.87
226 qFTA10&BnaC08cqDTF N78 × N133 38.20±2.49 cde 0.43 15.91 43.95
450 qFTA10&BnaC08cqDTF N80 × N133 34.00±0.71 hijk 0 0 39.97
451 qFTA10&BnaC08cqDTF N80 × N133 35.00±0.71 ghi 0 6.35 39.68
452 qFTA10&BnaC08cqDTF N80 × N133 35.40±0.55 gh 4.09 178.53 33.34
458 qFTA10&BnaC08cqDTF N80 × N133 34.60±0.55 hij 0.29 0 39.82

图4

含有双早花位点资源与单位点亲本花期对比图 图A从左向右依次是含有qFT.A02-1位点材料N103, qFT.A02-1&BnaC08cqDTF位点材料65, BnaC08cqDTF位点材料N133; 图B从左向右依次是含有BnaC08cqDTF位点材料N133, qFTA10&BnaC08cqDTF位点材料195, qFTA10位点材料N78; 图C从左向右依次是含有qFT.A02-1位点材料N103, qFT.A02-1&qFTA10位点材料12, qFTA10位点材料N78。"

表7

195连续2年测产统计"

年份
Year		 品种
Variety		 产量
Yield (kg)		 平均产量
Average yield (kg)		 比对照增产
Yield increase over control (%)
湟源
Huangyuan		 贵南
Guinan		 化隆
Hualong		 门源
Menyuan		 同德
Tongde		 海晏
Haiyan
2023 QH283 3275.00±417.58 b 1475.00±401.62 bc 5525.00±108.97 ab 2700.00±914.81 c 3400.00±86.60 c — 3275.00 ab −19.82
QH355 3733.33±1437.58 b 1318.75±249.30 c 5375.00±676.39 ab 3566.67±603.29 bc 2900.00±132.29 d — 3378.75 ab −17.28
浩油11号
Haoyou 11		 5375.00±1112.71 a 1750.00±307.21 ab 5741.67±387.57 ab 5183.33±289.76 a 2373.33±248.26 e — 4084.67 a 0
195 4683.33±700.59 ab 2231.25±92.07 a 5250.00±887.76 ab 4741.85±501.87 ab 4566.67±125.83 a — 4294.62 a 5.14
甘蓝型红花
Brassica napus with red flowers		 3191.67±687.54 b 1200.00±49.61 c 5800.00±640.80 a 2766.67±274.24 c 3833.33±76.38 b — 3358.33 ab −17.78
白菜型红花
Brassica rapa with red flowers		 3300.00±732.72 b 1237.50±99.22 c 4766.67±52.04 b 3275.00±325.00 c 2383.33±189.30 e — 2992.50 b −26.74
2024 QHYM05 — 2083.33±100.45 b 1612.50±141.97 d 2270.83±194.19 b 1610.42±275.02 b 812.50±225.09 ab 1677.92 c 10.33
QH241 — 2812.50±112.67 a 4114.58±90.21 a 3054.17±381.88 a 2291.67±570.68 a 747.92±139.8 abc 2604.17 a 71.23
浩油11号
Haoyou 11		 — 1968.75±360.39 b 1291.67±165.04 d 2270.83±194.19 b 1420.83±111.16 c 652.08±254.21 bcd 1520.83 c 0
195 — 2905.00±283.50 a 3424.95±452.34 b 3241.65±185.55 a 2452.95±395.70 a 996.05±144.15 a 2604.12 a 71.23
甘蓝型红花
Brassica napus with red flowers		 — 2197.92±47.74 b 2427.08±470.14 c 2433.33±301.13 b 1295.83±267.02 c 468.75±153.22 cd 1764.58 b 16.02
白菜型红花
Brassica rapa with red flowers		 — 1510.42±172.11 c 1283.33±189.71 d 2270.83±194.19 b 904.17±47.32 c 347.92±56.37 d 1263.33 d −16.93
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