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作物学报

作物学报 ›› 2019, Vol. 45 ›› Issue (12): 1806-1821.doi: 10.3724/SP.J.1006.2019.94027

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

甘蓝型油菜湿害胁迫响应性状的全基因组关联分析及候选基因预测

李阳阳1,2,3,荆蓉蓉1,3,吕蓉蓉1,2,3,石鹏程1,2,3,李欣1,2,3,王芹1,2,3,吴丹1,2,3,周清元1,3,李加纳1,2,3,唐章林1,2,3,*()   

  1. 1 西南大学农学与生物科技学院, 重庆 400715
    2 西南大学农业科学研究院, 重庆 400715
    3 重庆市油菜工程技术研究中心, 重庆 400715
  • 收稿日期:2019-02-25 接受日期:2019-06-24 出版日期:2019-12-12 网络出版日期:2019-07-16
  • 通讯作者: 唐章林
  • 作者简介:李阳阳, E-mail: liyangyangswu@163.com;|荆蓉蓉, E-mail: m15525440862@163.com
  • 基金资助:
    本研究由重庆市社会事业与民生保障科技创新主题专项(cstc2016shms-ztzx80010);国家重点研发计划项目(2018YFD0100504);中央高校基本科研业务费专项资金项目(XDJK2019D021)

Genome-wide association analysis and candidate genes prediction of waterlogging-responding traits in Brassica napus L.

Yang-Yang LI1,2,3,Rong-Rong JING1,3,Rong-Rong LYU1,2,3,Peng-Cheng SHI1,2,3,Xin LI1,2,3,Qin WANG1,2,3,Dan WU1,2,3,Qing-Yuan ZHOU1,3,Jia-Na LI1,2,3,Zhang-Lin TANG1,2,3,*()   

  1. 1 College of Agronomy and Biotechnology, Southwest University, Chongqing 400715, China
    2 Academy of Agricultural Sciences, Southwest University, Chongqing 400715, China
    3 Chongqing Engineering Research Center for Rapeseed, Chongqing 400715, China
  • Received:2019-02-25 Accepted:2019-06-24 Published:2019-12-12 Published online:2019-07-16
  • Contact: Zhang-Lin TANG
  • Supported by:
    This study was supported by the Science and Technology Innovation Project of Chongqing Social Undertakings & Livelihood Security(cstc2016shms-ztzx80010);the National Key R&D Program of China(2018YFD0100504);the Fundamental Research Funds for the Central Universities(XDJK2019D021)

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摘要：

湿害严重影响油菜的产量和品质, 前人在耐湿机制和生理响应等方面已经做了许多研究, 但涉及油菜耐湿相关基因的研究相对较少。本研究以248份甘蓝型油菜品种(系)为材料, 进行湿害胁迫处理, 调查了8个湿害响应的相关性状, 基于60K Illumina Infinium SNP芯片基因型数据对各性状耐湿系数进行了全基因组关联分析和湿害响应候选基因预测。结果显示, 湿害胁迫较正常灌溉油菜幼苗绿叶数减少, 地上部干重和鲜重以及叶片可溶性蛋白含量降低, 叶片POD酶活性和MDA含量略有升高。地下部干重和鲜重在多数材料中降低, 而在有些材料中升高, 变化幅度均较小。经全基因组关联分析共检测到与耐湿系数显著关联的SNP标记36个, 可解释表型变异8.28%~12.95%, 其中17个所关联的耐湿系数在基因型间具有显著(P<0.05)或极显著(P<0.01)差异, 其所在的Blocks共覆盖了71个候选基因, 包括Blast到同源拟南芥基因64个, 主要编码转录因子(如Transcription initiation factor IIF、bZIP68、myb-like HTH transcriptional regulator family protein)、转运蛋白(如SUC2)、生长抑制子(如ING2)、蛋白磷酸酶(如Protein phosphatase 2C family protein)、DNA/RNA结合蛋白[如bHLH DNA-binding superfamily protein、RNA-binding (RRM/RBD/RNP motifs) family protein]、激素响应蛋白(如ARF8、ADC2、ERD6、NF-YC9)以及氧化胁迫、渗透胁迫、盐胁迫或水分剥夺响应蛋白(如ERD6、NP1、TIR-NBS-LRR、CRT1b)等。本研究可为揭示甘蓝型油菜耐湿机理和培育耐湿新品种奠定基础。

关键词: 甘蓝型油菜, 湿害胁迫响应性状, 全基因组关联分析, 候选基因

Abstract:

The yield and quality of rapeseed are significantly affected by waterlogging. Many researches on waterlogging of rapeseed mostly aimed at its mechanism and physiological response, but a few of them identified the genes related to waterlogging. In this study, 248 Brassica napus L. accessions were planted in flowerpots with treatments of waterlogging and well watering for four weeks when the plants had four true leaves in rain-proof installation, and eight waterlogging-responding traits were investigated. The genome-wide association analysis was carried out based on 60K Illumina Infinium SNP genotype data and the candidate genes associated with waterlogging responses were predicted. Under waterlogging, the number of green leaf, the shoot fresh weight and dry weight, the leaf soluble protein content were lower, and the leaf POD activity, the MDA content were higher than those under well watering. The root fresh weight and dry weight were lower in most accessions and higher in some accessions under waterlogging stress. A total of 36 SNPs were identified associated with waterlogging traits and explained the phenotypic variation of 8.28%-12.95%. There were significant (P < 0.05) or extremely significant (P < 0.01) differences in the waterlogging resistance index among genotypes of 17 SNPs. The Blocks containing 17 SNPs covered 71 candidate genes. Among them, 64 candidate genes were homologous to Arabidopsis genes by blast. They encoded transcription factors (e.g. transcription initiation factor IIF, bZIP68, myb-like HTH transcriptional regulator family protein), transport protein (e.g. SUC2), inhibitor of growth (e.g. ING2), protein phosphatase (e.g. protein phosphatase 2C family protein), DNA/RNA binding proteins [e.g. bHLH DNA-binding superfamily protein, RNA-binding (RRM/RBD/RNP motifs) family protein], hormone response proteins (e.g. ARF8, ADC2, ERD6, NF-YC9), oxidative/osmotic/cold stress or water deprivation response proteins (e.g. ERD6, NP1, TIR-NBS-LRR, CRT1b). These results would be benefit for revealing the waterlogging resistance mechanism and cultivating new varieties with waterlogging resistance in Brassica napus L.

Key words: Brassica napus L., waterlogging-responding traits, genome-wide association analysis, candidate genes

图1

甘蓝型油菜幼苗湿害胁迫响应性状耐湿系数频数分布图 缩写同表1。"

表1

甘蓝型油菜幼苗湿害胁迫响应性状的方差分析"

变异来源
Source of variation		 自由度
df		 F-value
GLN SFW SDW RFW RDW POD MDA Protein
水分间Water 1 4840.51** 29,719.19** 6028.69** 67.37** 1113.40** 1167.63** 2314.55** 17,856.96**
材料间Variety 245 7.89** 38.19** 17.30** 11.25** 13.21** 12.81** 34.72** 20.95**
水分×材料Water × Variety 245 1.95** 20.63** 8.97** 8.08** 7.62** 13.27** 31.70** 17.80**
误差Error 984

图2

248份甘蓝型油菜品种(系)的亲缘系数频率分布图"

图3

2个连续K值所对应的后验概率的变化速率(ΔK)"

图4

248份甘蓝型油菜品种(系)群体结构分布图"

图5

甘蓝型油菜染色体连锁不平衡衰减曲线"

表2

甘蓝型油菜各染色体的衰减距离"

染色体Chromosome 衰减距离
LD decay (kb)		 染色体Chromosome 衰减距离
LD decay (kb)
A01 180 C01 750
A02 180 C02 1200
A03 180 C03 400
A04 180 C04 590
A05 180 C05 290
A06 180 C06 400
A07 180 C07 380
A08 500 C08 620
A09 380 C09 710
A10 210

图6

甘蓝型油菜湿害胁迫响应性状耐湿系数基于6种模型的关联分析QQ图"

图7

甘蓝型油菜湿害胁迫响应性状耐湿系数基于最佳模型关联分析的Manhattan图"

表3

甘蓝型油菜湿害胁迫响应性状WLRI显著关联的SNP标记"

性状
Trait		 标记
Marker		 染色体
Chromosome		 位置
Position		 P值
P-value		 贡献率
R2 (%)
GLN Bn-scaff_23954_1-p1045541 C03 10,880,567 1.61E-05 9.82
Bn-scaff_20901_1-p1110262 C05 2,905,509 5.17E-06 10.69
Bn-A03-p11122273 C07 41,947,271 5.09E-06 10.70
性状
Trait		 标记
Marker		 染色体
Chromosome		 位置
Position		 P值
P-value		 贡献率
R2 (%)
SFW Bn-A02-p13803435 A02 10,432,008 1.36E-05 9.71
Bn-A04-p6975646a A04 8,262,555 8.53E-06 10.06
Bn-A04-p6978163b A04 8,265,085 8.53E-06 10.06
Bn-A04-p6980459 A04 8,267,518 1.64E-05 9.57
Bn-A10-p15740896c A10 15,809,264 1.46E-05 9.65
Bn-scaff_16531_1-p428890d C04 21,584,887 7.76E-06 10.13
SDW Bn-A04-p6975646a A04 8,262,555 1.71E-05 9.74
Bn-A04-p6978163b A04 8,265,085 1.71E-05 9.74
Bn-A10-p15740896c A10 15,809,264 1.94E-06 11.40
Bn-scaff_21884_1-p644186 C01 37,469,955 7.66E-06 10.36
Bn-scaff_16531_1-p428890d C04 21,584,887 4.67E-06 10.73
Bn-scaff_17919_1-p94976 C09 34,508,780 2.82E-05 8.28
RFW Bn-scaff_18062_1-p214669 C04 31,128,751 2.45E-05 9.47
Bn-scaff_21124_1-p301252 C05 37,592,772 7.89E-06 10.35
RDW Bn-A08-p15326882 A08 12,894,573 2.74E-06 12.39
Bn-A09-p20263302 A09 17,272,800 1.95E-06 12.71
Bn-scaff_18826_1-p776957 C05 36,457,927 1.02E-05 11.14
Bn-scaff_17888_1-p170198 C09 15,826,668 1.74E-05 10.64
Protein Bn-A01-p6538362 A01 6,002,545 2.06E-05 10.54
Bn-A02-p12822311 A02 9,572,851 8.39E-06 11.47
Bn-A03-p7079712 A03 6,365,301 2.87E-05 11.61
Bn-A03-p20100490 A03 18967298 1.57E-05 10.83
Bn-A07-p20887568 A07 22,470,590 7.18E-06 11.63
Bn-A09-p21935602 A09 19,325,386 2.67E-06 12.65
Bn-scaff_20675_1-p32364 C01 9,755,072 2.08E-05 11.96
Bn-scaff_21188_1-p1142 C02 17,317,511 8.30E-06 12.95
Bn-scaff_17067_1-p509745 C02 27,930,748 2.18E-05 11.91
Bn-scaff_15585_1-p1047449 C04 44,459,141 2.45E-06 12.74
POD Bn-A01-p637586 A01 228,343 4.62E-06 9.61
Bn-A03-p12250401 A03 11,367,398 1.93E-05 9.62
Bn-A09-p30788157 A09 28,540,524 7.44E-06 10.35
Bn-A10-p2246688 A10 1,588,516 2.00E-05 9.59
MDA Bn-A06-p3088460 C05 3,221,158 2.31E-05 9.38

表4

甘蓝型油菜湿害胁迫响应性状显著关联SNP标记不同基因型的耐湿系数"

性状
Trait		 SNP标记
SNP marker		 基因型
Loci		 耐湿系数
WLRI		 方差
Variance
GLN Bn-scaff_23954_1-p1045541 (GG)5 0.793 Aa 0.005
(TG)25 0.627 Bb 0.012
(TT)168 0.589 Bb 0.008
Bn-scaff_20901_1-p1110262 (AA)50 0.656 Aa 0.013
(CC)155 0.593 Bb 0.007
(AC)20 0.570 Bb 0.009
Bn-A03-p11122273 (CC)25 0.681 Aa 0.010
(TC)5 0.647 ABab 0.007
(TT)202 0.586 Bb 0.008
性状
Trait		 SNP标记
SNP marker		 基因型
Loci		 耐湿系数
WLRI		 方差
Variance
SFW Bn-A02-p13803435 (TT)210 0.361 Aa 0.022
(TC)12 0.347 ABab 0.016
(CC)16 0.296 Bb 0.006
Bn-A04-p6975646 (AA)12 0.560 Aa 0.036
(CC)220 0.355 Bb 0.020
(AC)8 0.273 Bb 0.017
Bn-A04-p6978163 (TT)12 0.560 Aa 0.036
(GG)222 0.356 Bb 0.021
(TG)8 0.273 Bb 0.017
Bn-A04-p6980459 (GG)13 0.570 Aa 0.034
AA)218 0.355 Bb 0.021
Bn-A10-p15740896 (TT)14 0.546 Aa 0.039
(TC)3 0.435 ABab 0.026
(CC)216 0.357 Bb 0.021
Bn-scaff_16531_1-p428890 (CC)4 0.767 Aa 0.022
(TC)17 0.380 Bb 0.014
(TT)200 0.358 Bb 0.023
SDW Bn-A04-p6975646 (AA)12 0.862 Aa 0.178
(CC)220 0.548 ABb 0.046
(AC)8 0.416 Bc 0.016
Bn-A04-p6978163 (TT)12 0.862 Aa 0.178
(GG)222 0.550 ABb 0.047
(TG)8 0.416 Bc 0.016
Bn-A10-p15740896 TT)14 0.865 Aa 0.098
(TC)3 0.756 ABab 0.170
(CC)216 0.544 Bb 0.048
Bn-scaff_21884_1-p644186 (GG)4 1.125 Aa 0.094
(AA)174 0.566 Bb 0.053
(AG)25 0.512 Bb 0.050
Bn-scaff_16531_1-p428890 (CC)4 1.145 Aa 0.138
(TC)17 0.618 Bb 0.061
(TT)200 0.545 Bb 0.049
Bn-scaff_17919_1-p94976 (AA)213 0.559 Aa 0.051
(AC)23 0.455 Bb 0.020
RFW Bn-scaff_18062_1-p214669 (AC)11 2.538 Aa 2.529
(CC)67 1.222 Ab 0.347
(AA)143 1.291 Ab 0.850
Bn-scaff_21124_1-p301252 (TT)208 1.363 Aa 0.969
(CC)28 1.317 Aa 0.697
(TC)6 0.988 Aa 0.132
RDW Bn-A08-p15326882 (AA)215 1.464 Aa 0.365
(AG)27 1.321 Aa 0.241
Bn-A09-p20263302 (TT)198 1.471 Aa 0.349
(TA)19 1.414 Aab 0.412
性状
Trait		 SNP标记
SNP marker		 基因型
Loci		 耐湿系数
WLRI		 方差
Variance
(AA)24 1.195 Ab 0.217
Bn-scaff_18826_1-p776957 (TC)26 1.546 Aa 0.318
(CC)69 1.469 Aa 0.334
(TT)136 1.367 Aa 0.317
Bn-scaff_17888_1-p170198 (AC)30 1.534 Aa 0.321
(CC)77 1.505 Aa 0.423
(AA)124 1.343 Aa 0.232
Protein Bn-A01-p6538362 (TT)117 0.293 Aa 0.025
(TG)108 0.291 Aa 0.023
Bn-A02-p12822311 (AA)174 0.302 Aa 0.028
(AG)67 0.302 Aa 0.024
Bn-A03-p7079712 (AG)7 0.681 Aa 0.122
(AA)33 0.312 Ab 0.034
(GG)197 0.295 Ab 0.024
Bn-A03-p20100490 (TC)50 0.318 Aa 0.029
(CC)189 0.296 Aa 0.025
Bn-A07-p20887568 (TC)75 0.299 Aa 0.023
(TT)162 0.298 Aa 0.027
Bn-A09-p21935602 (GG)217 0.314 Aa 0.031
(AG)26 0.230 Bb 0.008
Bn-scaff_20675_1-p32364 (GG)9 0.358 Aa 0.039
(TG)14 0.305 Aa 0.028
(TT)208 0.291 Aa 0.023
Bn-scaff_21188_1-p1142 (GG)3 0.350 Aa 0.046
(TT)193 0.293 Aa 0.022
(TG)33 0.282 Aa 0.033
Bn-scaff_17067_1-p509745 (TG)16 0.330 Aa 0.054
(TT)33 0.313 Aa 0.029
(GG)191 0.297 Aa 0.024
Bn-scaff_15585_1-p1047449 (GG)211 0.301 Aa 0.026
(AG)28 0.286 Aa 0.030
POD Bn-A01-p637586 (AA)212 1.195 Aa 0.089
(AG)28 1.118 Aa 0.066
Bn-A03-p12250401 (AA)40 1.188 Aa 0.086
(GG)162 1.186 Aa 0.076
(AG)34 1.150 Aa 0.121
Bn-A09-p30788157 (TT)225 1.201 Aa 0.084
(CC)13 0.980 Bb 0.059
Bn-A10-p2246688 (CC)34 1.282 Aa 0.115
(AA)182 1.168 Aa 0.081
(AC)20 1.146 Aa 0.092
MDA Bn-A06-p3088460 (GG)195 1.513 Aa 0.489
(AG)47 1.294 Ab 0.403

图8

显著性标记所在的Blocks及候选基因 显著性标记用黑色边框标示, 候选基因用下画线标示。"

表5

甘蓝型油菜湿害胁迫响应性状耐湿系数候选基因相关信息"

油菜基因
B. napus gene		 染色体
Chr.		 物理位置
Position (bp)		 拟南芥基因
AT gene		 功能
Function
BnaA02g17270D A02 10,385,018-10,386,352 AT1G75510 Transcription initiation factor IIF
BnaA02g17280D A02 10,389,281-10,390,641 AT1G75520 SHI-related sequence 5 (SRS5)
BnaA02g17290D A02 10,428,468-10,429,854 AT1G75540 Salt tolerance homolog 2 (STH2)
BnaA02g17300D A02 10,461,565-10,461,997 AT1G75580 SAUR-like auxin-responsive protein family
BnaA03g13900D A03 6,364,664-6,366,026 AT2G30360 SOS3-interacting protein 4 (SIP4)
BnaA04g09140D A04 8,212,219-8,213,486 AT5G39320 UDP-glucose 6-dehydrogenase family protein
BnaA04g09150D A04 8,215,808-8,217,193 AT5G39050 HXXXD-type acyl-transferase family protein
BnaA04g09160D A04 8,231,509-8,231,884 AT1G22710 ARABIDOPSIS THALIANA SUCROSE-PROTON SYMPORTER 2 (SUC2)
BnaA04g09170D A04 8,231,950-8,233,460 AT5G39340 Histidine-containing phosphotransmitter 3 (AHP3)
BnaA04g09180D A04 8,233,647-8,235,665 AT5G39350 Tetratricopeptide repeat (TPR)-like superfamily protein
BnaA04g09190D A04 8,235,944-8,237,287 AT5G39360 EID1-like 2 (EDL2)
BnaA04g09200D A04 8,238,128-8,241,036 AT5G39380 Plant calmodulin-binding protein-related
BnaA04g09210D A04 8,261,813-8,262,055 —
BnaA04g09220D A04 8,280,136-8,282,045 AT5G39400 PTEN1
BnaA04g09230D A04 8,282,166-8,284,176 AT5G39410 Saccharopine dehydrogenase
BnaA09g24570D A09 17,276,994-17,280,062 AT1G32190 Alpha/beta-Hydrolases superfamily protein
BnaA09g24580D A09 17,282,321-17,283,676 —
BnaA09g24590D A09 17,303,368-17,305,678 AT1G46984 F-box family protein
BnaA09g24600D A09 17,320,367-17,322,665 AT1G46984 F-box family protein
BnaA09g24610D A09 17,360,901-17,363,794 AT1G32150 Basic region/leucine zipper transcription factor 68 (bZIP68)
BnaA09g26170D A09 19,351,820-19,355,843 AT5G37020 Auxin response factor 8 (ARF8)
BnaA09g40620D A09 28,538,438-28,542,878 AT2G26260 3 beta-hydroxysteroid-dehydrogenase/decarboxylase isoform 2 (3BETAHSD/D2)
BnaA10g24140D A10 15,809,477-15,810,280 AT4G34710 Arginine decarboxylase 2 (ADC2)
BnaA10g24150D A10 15,810,902-15,813,379 AT5G06800 Myb-like HTH transcriptional regulator family protein
BnaC03g20520D C03 10,879,272-10,881,026 AT2G37110 PLAC8 family protein
BnaC04g20480D C04 21,584,060-21,585,266 AT1G63740 Disease resistance protein (TIR-NBS-LRR class) family
BnaC04g29520D C04 31,077,933-31,080,151 AT4G12790 P-loop containing nucleoside triphosphate hydrolases superfamily protein
BnaC04g29530D C04 31,085,683-31,088,293 AT4G12740 HhH-GPD base excision DNA repair family protein
油菜基因
B. napus gene		 染色体
Chr.		 物理位置
Position (bp)		 拟南芥基因
AT gene		 功能
Function
BnaC04g29540D C04 31,096,099-31,098,880 AT4G12650 Endomembrane protein 70 protein family
BnaC04g29550D C04 31,113,638-31,113,859 —
BnaC04g29560D C04 31,116,424-31,116,591 —
BnaC04g29570D C04 31,121,991-31,124,968 AT4G12680 Unknown protein
BnaC04g29580D C04 31,130,755-31,131,747 AT1G54390 INHIBITOR OF GROWTH 2 (ING2)
BnaC04g29590D C04 31,133,548-31,136,282 AT5G35380 Protein kinase protein with adenine nucleotide alpha hydrolases-like domain
BnaC04g29600D C04 31,158,769-31,162,504 AT5G35430 Tetratricopeptide repeat (TPR)-like superfamily protein
BnaC04g29610D C04 31,170,772-31,172,233 AT5G35520 MINICHROMOSOME INSTABILITY 12 (MIS12)-LIKE (MIS12)
BnaC05g05880D C05 2,904,698-2,910,332 AT1G08190 Vacuolar protein sorting 41 (VPS41)
BnaC05g06500D C05 3,218,324-3,221,856 AT1G08930 EARLY RESPONSE TO DEHYDRATION 6 (ERD6)
BnaC05g06510D C05 3,222,332-3,224,004 AT1G08940 Phosphoglycerate mutase family protein
BnaC05g06520D C05 3,224,198-3,228,539 AT1G08970 Nuclear factor Y
BnaC05g06530D C05 3,228,942-3,233,245 AT1G09000 NPK1-related protein kinase 1 (NP1)
BnaC05g06540D C05 3,233,325-3,237,550 AT1G09010 Glycoside hydrolase family 2 protein
BnaC05g06550D C05 3,239,409-3,244,162 AT1G09020 Homolog of yeast sucrose nonfermenting 4 (SNF4)
BnaC05g06560D C05 3,246,008-3,247,080 —
BnaC05g06570D C05 3,248,229-3,248,627 AT1G09030 Nuclear factor Y
BnaC05g06580D C05 3,258,505-3,259,753 AT2G19610 RING/U-box superfamily protein
BnaC05g06590D C05 3,261,801-3,265,778 AT1G09050 Unknown protein
BnaC05g06600D C05 3,266,250-3,266,895 AT1G09060 Zinc finger protein
BnaC05g06610D C05 3,267,487-3,268,443 AT1G09070 Soybean gene regulated by cold-2 (SRC2)
BnaC05g06620D C05 3,269,846-3,271,741 AT1G09130 ATP-dependent caseinolytic (Clp) protease/crotonase family protein
BnaC05g06630D C05 3,275,956-3,279,260 AT1G09140 SERINE-ARGININE PROTEIN 30 (ATSRP30)
BnaC05g06640D C05 3,279,332-3,280,873 AT1G09150 Containing protein
BnaC05g06650D C05 3,281,248-3,282,283 AT1G09155 Phloem protein 2-B15 (PP2-B15)
BnaC05g06660D C05 3,298,672-3,298,972 AT1G09155 Phloem protein 2-B15 (PP2-B15)
BnaC05g06670D C05 3,299,052-3,303,054 AT1G09155 Phloem protein 2-B15 (PP2-B15)
BnaC05g06680D C05 3,303,126-3,306,459 AT2G43445 F-box and associated interaction domains-containing protein
BnaC05g06690D C05 3,308,135-3,309,772 AT1G09160 Protein phosphatase 2C family protein
BnaC05g06700D C05 3,311,000-3,312,454 AT1G09190 Tetratricopeptide repeat (TPR)-like superfamily protein
BnaC05g06710D C05 3,312,506-3,313,708 AT1G56330 Secretion-associated RAS 1B (SAR1B)
BnaC05g06720D C05 3,316,670-3,319,893 AT5G08020 RPA70-kDa subunit B (RPA70B)
BnaC05g06730D C05 3,321,779-3,322,096 —
BnaC05g06740D C05 3,324,873-3,325,493 AT5G09390 CD2-binding protein-related
BnaC05g06750D C05 3,329,829-3,330,444 AT1G09250 Basic helix-loop-helix (bHLH) DNA-binding superfamily protein
BnaC05g06760D C05 3,330,666-3,331,280 AT1G09250 Basic helix-loop-helix (bHLH) DNA-binding superfamily protein
BnaC05g06770D C05 3,349,687-3,350,292 —
BnaC05g06780D C05 3,356,126-3,359,392 AT1G09210 Calreticulin 1b (CRT1b)
BnaC05g06790D C05 3,360,115-3,363,017 AT1G09230 RNA-binding (RRM/RBD/RNP motifs) family protein
BnaC05g06800D C05 3,370,500-3,371,711 AT1G09240 Nicotianamine synthase 3 (NAS3)
BnaC05g06810D C05 3,372,700-3,373,315 AT1G09250 Basic helix-loop-helix (bHLH) DNA-binding superfamily protein
BnaC05g06820D C05 3,378,631-3,379,149 AT1G09260 Chaperone DnaJ-domain superfamily protein
BnaC05g06830D C05 3,379,950-3,383,047 AT1G09270 Importin alpha isoform 4 (IMPA-4)
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