苹果GRAS全基因组鉴定及其对生长素的响应分析

doi:10.16420/j.issn.0513-353x.2021-1039

摘要/Abstract

摘要：

从苹果（Malus × domestica）全基因组中鉴定到78个结构完整的GRAS基因，它们的编码蛋白分属于11个亚家族，其中LISCL亚家族成员最多；这78个MdGRAS基因分别定位于18条染色体上，其中11个位于3号染色体上。苹果GRAS家族中还存在片段复制和串联复制基因。苹果器官特异性表达谱显示，除MdGRAS47外，大部分MdGRAS在根系中表达量较低；MdGRAS20在枝条中表达量较高，MdGRAS18、MdGRAS55和MdGRAS61在种子中表达量较高。启动子分析显示，这78个MdGRAS全部含光响应元件，76个含胁迫相关元件，37个含有生长素响应元件；17个MdGRAS的表达对2,4-D处理有响应，其中MdGRAS30和MdGRAS61的响应最显著。

关键词: 苹果, GRAS基因家族, 生物信息, 启动子, 生长素

Abstract:

GRAS is a plant-specific transcription factor. In this study，78 GRAS genes were identified from the whole genome of apple（Malus × domestica）based on the conserved domain PF0351. MdGRASs encoding proteins belong to 11 subfamilies and most of them are the members of LISCL subfamily. They are located on 18 chromosomes，11 of which are located on chromosome 3. There were segmental duplication and tandem duplication events in GRAS family of apple. Ogran-specific expression profile analysis showed that，except for MdGRAS47，most of MdGRASs exhibited low transcript abundance levels in apple roots；MdGRAS20 showed higher expression level in shoots while MdGRAS18，MdGRAS55 and MdGRAS61 highly expressed in seeds. Promoter analysis indicated that all 78 MdGRASs contained light response elements，76 out of 78 MdGRASs contained stress-related elements and 37 out of 78 MdGRASs contained auxin response elements. It was confirmed that 17 MdGRASs could respond to 2,4-D with MdGRAS30 and MdGRAS61 responded to 2,4-D most significantly.

Key words: apple, GRAS gene family, bioinformatics, promoter, auxin

中图分类号:

S661.1

于婷婷, 李欢, 宁源生, 宋建飞, 彭璐琳, 贾竣淇, 张玮玮, 杨洪强. 苹果GRAS全基因组鉴定及其对生长素的响应分析[J]. 园艺学报, 2023, 50(2): 397-409.

YU Tingting, LI Huan, NING Yuansheng, SONG Jianfei, PENG Lulin, JIA Junqi, ZHANG Weiwei, YANG Hongqiang. Genome-wide Identification of GRAS Gene Family in Apple and Expression Analysis of Its Response to Auxin[J]. Acta Horticulturae Sinica, 2023, 50(2): 397-409.

图/表 6

参考文献 35

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名称 Primer	碱基序列（5′-3′） Sequence	名称 Primer	碱基序列（5′-3′） Sequence
MdGRAS8-F	ACAGAGCAAGAGGCAAACCA	MdGRAS8-R	CTCTCCGTTCGCTCAGTTCC
MdGRAS13-F	GAGGGTAAGCCCTGCATGTT	MdGRAS13-R	CCCCAGTCTGTCTTGGTAGC
MdGRAS14-F	TGATCCTCAATGGGTTGGCG	MdGRAS14-R	AAGAGCCCTGCAGCACATTA
MdGRAS30-F	CGGTTGTGGATAGGTTGGAG	MdGRAS30-R	ATTGTGCTCAGCCTCTTGCT
MdGRAS34-F	AGGGATCGCGTGGAAAGATG	MdGRAS34-R	AGAGTGCGGCTTATCCAACC
MdGRAS41-F	TCCAACTCGTCCACACACTC	MdGRAS41-R	GGAAGAATCGAGGCAATCCT
MdGRAS44-F	TCACTTCCAACCAAGCCATC	MdGRAS44-R	TGAGAGTTGCTTTCCGGTCT
MdGRAS50-F	TTCAAACAGGCGCTCTTCCA	MdGRAS50-R	CCGATCCTCGCATACCTAGC
MdGRAS53-F	TGACTCCCTAGATGCCATGC	MdGRAS53-R	CCTGCGCCAGCATCTTAATC
MdGRAS55-F	AGATGCAGAGGTCCATGTCA	MdGRAS55-R	GAGAGCTTCAGCACATGCAA
MdGRAS56-F	GCAGCGCCTTGATGATATGC	MdGRAS56-R	CCATCGGCACACTTGCTTTC
MdGRAS61-F	CAGCAACACTTACTGCACCC	MdGRAS61-R	TGAAAGGCACCCCCATCAAC
MdGRAS62-F	ACCGGAATTCCCCAATCGAG	MdGRAS62-R	CATCTCCGCCAACGTCTTCT
MdGRAS63-F	GGAGGGTGCTAAATGGGCTT	MdGRAS63-R	GGTTTGTGCAAAACTCCCCA
MdGRAS68-F	GCTCTTCAGTCCCTCCTCAA	MdGRAS68-R	ATGTCGAAGTCGATGACGTG
MdGRAS69-F	CAGCAACACTTACTGCACCC	MdGRAS69-R	TGAAAGGCACCCCCATCAAC
MdGRAS75-F	GGAGGGTGCTAAATGGGCTT	MdGRAS75-R	TGTAGCACCGGAGACCTGAT

名称 Primer	碱基序列（5′-3′） Sequence	名称 Primer	碱基序列（5′-3′） Sequence
MdGRAS8-F	ACAGAGCAAGAGGCAAACCA	MdGRAS8-R	CTCTCCGTTCGCTCAGTTCC
MdGRAS13-F	GAGGGTAAGCCCTGCATGTT	MdGRAS13-R	CCCCAGTCTGTCTTGGTAGC
MdGRAS14-F	TGATCCTCAATGGGTTGGCG	MdGRAS14-R	AAGAGCCCTGCAGCACATTA
MdGRAS30-F	CGGTTGTGGATAGGTTGGAG	MdGRAS30-R	ATTGTGCTCAGCCTCTTGCT
MdGRAS34-F	AGGGATCGCGTGGAAAGATG	MdGRAS34-R	AGAGTGCGGCTTATCCAACC
MdGRAS41-F	TCCAACTCGTCCACACACTC	MdGRAS41-R	GGAAGAATCGAGGCAATCCT
MdGRAS44-F	TCACTTCCAACCAAGCCATC	MdGRAS44-R	TGAGAGTTGCTTTCCGGTCT
MdGRAS50-F	TTCAAACAGGCGCTCTTCCA	MdGRAS50-R	CCGATCCTCGCATACCTAGC
MdGRAS53-F	TGACTCCCTAGATGCCATGC	MdGRAS53-R	CCTGCGCCAGCATCTTAATC
MdGRAS55-F	AGATGCAGAGGTCCATGTCA	MdGRAS55-R	GAGAGCTTCAGCACATGCAA
MdGRAS56-F	GCAGCGCCTTGATGATATGC	MdGRAS56-R	CCATCGGCACACTTGCTTTC
MdGRAS61-F	CAGCAACACTTACTGCACCC	MdGRAS61-R	TGAAAGGCACCCCCATCAAC
MdGRAS62-F	ACCGGAATTCCCCAATCGAG	MdGRAS62-R	CATCTCCGCCAACGTCTTCT
MdGRAS63-F	GGAGGGTGCTAAATGGGCTT	MdGRAS63-R	GGTTTGTGCAAAACTCCCCA
MdGRAS68-F	GCTCTTCAGTCCCTCCTCAA	MdGRAS68-R	ATGTCGAAGTCGATGACGTG
MdGRAS69-F	CAGCAACACTTACTGCACCC	MdGRAS69-R	TGAAAGGCACCCCCATCAAC
MdGRAS75-F	GGAGGGTGCTAAATGGGCTT	MdGRAS75-R	TGTAGCACCGGAGACCTGAT

基因名 Gene name	基因ID Gene ID	起始位点 Start site	终止位点 End site	染色体 Chromosome	CDS/bp	蛋白质性质Deduce polypetide
基因名 Gene name	基因ID Gene ID	起始位点 Start site	终止位点 End site	染色体 Chromosome	CDS/bp	长度/aa Length	分子量/Da MW	等电点 pI
MdGRAS1	MD00G1043300	7 80 6171	7 809 643	Chrom00	2 424	808	88 359.54	5.80
MdGRAS2	MD00G1112400	23 667 785	23 669 215	Chrom00	1 428	476	53 228.76	5.67
MdGRAS3	MD01G1206800	30 231 215	30 233 158	Chrom01	1 941	647	70 816.40	6.63
MdGRAS4	MD02G1039600	3 216 839	3 218 479	Chrom02	1 638	546	59 826.80	5.63
MdGRAS5	MD02G1148800	12 235 195	12 237 263	Chrom02	1 812	604	66 498.43	5.17
MdGRAS6	MD02G1185900	16 892 351	16 893 874	Chrom02	1 521	507	56 882.60	5.22
MdGRAS7	MD02G1210200	21 832 327	21 834 579	Chrom02	2 250	750	81 783.01	5.80
MdGRAS8	MD02G1295800	35 009 786	35 012 501	Chrom02	1 374	458	51 566.29	5.67
MdGRAS9	MD03G1045700	3 674 587	3 675 963	Chrom03	1 374	458	52 030.80	6.04
MdGRAS10	MD03G1088900	7 330 830	7 332 881	Chrom03	2 049	683	77 700.00	5.52
MdGRAS11	MD03G1089100	7 337 571	7 339 625	Chrom03	2 052	684	78 076.35	6.42
MdGRAS12	MD03G1089200	7 340 949	7 345 605	Chrom03	3 828	1 276	144 938.23	5.93
MdGRAS13	MD03G1089300	7 346 485	7 348 548	Chrom03	2 061	687	77 570.95	5.49
MdGRAS14	MD03G1089700	7 367 711	7 371 218	Chrom03	2 427	809	91 167.55	5.41
MdGRAS15	MD03G1089800	7 380 517	7 382 781	Chrom03	2 208	736	82 619.61	5.28
MdGRAS16	MD03G1169300	22 929 145	22 930 705	Chrom03	738	246	27 338.81	5.90
MdGRAS17	MD03G1188800	25 920 210	25 922 997	Chrom03	1 611	537	59 547.07	5.15
MdGRAS18	MD03G1214700	29 482 170	29 485 609	Chrom03	1 749	583	64 679.63	5.73
MdGRAS19	MD03G1293700	37 023 448	37 025 172	Chrom03	1 722	574	65 184.13	5.94
MdGRAS20	MD04G1046000	5 332 407	5 334 295	Chrom04	1 278	426	47 811.20	5.21
MdGRAS21	MD04G1088100	13 542 153	13 544 592	Chrom04	2 382	794	86 879.60	6.00
MdGRAS22	MD04G1163300	25 473 582	25 475 192	Chrom04	1 608	536	60 735.28	4.67
MdGRAS23	MD04G1163400	25 477 063	25 478 646	Chrom04	1 581	527	59 593.09	4.82
MdGRAS24	MD04G1181500	27 256 030	27 259 385	Chrom04	2 487	829	88 693.53	5.88
MdGRAS25	MD05G1011200	2 343 179	2 345 898	Chrom05	1 737	579	64 386.15	4.96
MdGRAS26	MD05G1040300	6 871 948	6 873 466	Chrom05	1 440	480	54 685.42	6.91
MdGRAS27	MD05G1194300	32 192 340	32 195 288	Chrom05	1 704	568	63 969.95	6.42
MdGRAS28	MD05G1229500	36 294 983	36 296 611	Chrom05	1 626	542	60 679.72	4.86
MdGRAS29	MD05G1360300	47 489 161	47 492 239	Chrom05	1 608	536	60 231.21	5.68
MdGRAS30	MD06G1073500	18 288 623	18 290 677	Chrom06	2 052	684	75 422.74	5.56
MdGRAS31	MD06G1148900	29 171 471	29 173 054	Chrom06	1 581	527	58 725.06	6.37
MdGRAS32	MD07G1030000	2 514 355	2 517 053	Chrom07	1 374	458	51 767.52	5.86
MdGRAS33	MD07G1115800	13 984 309	13 986 546	Chrom07	2 235	745	81 207.10	5.50
MdGRAS34	MD07G1277400	34 273 655	34 275 613	Chrom07	1 956	652	71 075.63	6.27
MdGRAS35	MD08G1167600	19 831 740	19 833 281	Chrom08	1 539	513	56 341.82	5.36
MdGRAS36	MD09G1007200	510 029	513 274	Chrom09	2 112	704	78 249.07	5.82
MdGRAS37	MD09G1103000	7 539 020	7 540 351	Chrom09	1 329	443	49 061.78	5.19
基因名 Gene name	基因ID Gene ID	起始位点 Start site	终止位点 End site	染色体 Chromosome	CDS/bp	蛋白质性质Deduce polypetide
基因名 Gene name	基因ID Gene ID	起始位点 Start site	终止位点 End site	染色体 Chromosome	CDS/bp	长度/aa Length	分子量/Da MW	等电点 pI
MdGRAS38	MD09G1121700	9 383 149	9 386 262	Chrom09	1 659	553	62 323.66	5.74
MdGRAS39	MD09G1137800	10 707 681	10 709 030	Chrom09	1 347	449	48 949.18	5.35
MdGRAS40	MD09G1183000	15 735 881	15 737 539	Chrom09	1 656	552	61 147.14	6.07
MdGRAS41	MD09G1264800	33 799 664	33 801 406	Chrom09	1 740	580	63 166.16	5.09
MdGRAS42	MD10G1011000	1 547 558	1 550 844	Chrom10	1 737	579	64 463.19	5.03
MdGRAS43	MD10G1046300	6 180 700	6 182 115	Chrom10	1 413	471	53 926.17	6.31
MdGRAS44	MD10G1181000	27 411 839	27 413 593	Chrom10	1 542	514	58 030.64	5.91
MdGRAS45	MD10G1336100	41 266 090	41 268 919	Chrom10	1 374	458	51 746.93	6.36
MdGRAS46	MD11G1047900	4 170 134	4 171 525	Chrom11	1 389	463	52 716.63	6.07
MdGRAS47	MD11G1097900	8 118 511	8 120 535	Chrom11	2 022	674	76 651.13	5.45
MdGRAS48	MD11G1098000	8 127 739	8 129 778	Chrom11	2 037	679	77 294.04	5.91
MdGRAS49	MD11G1098100	8 130 580	8 133 055	Chrom11	2 016	672	76 660.73	6.37
MdGRAS50	MD11G1098400	8 139 575	8 141 815	Chrom11	2 166	722	81 747.16	5.92
MdGRAS51	MD11G1099000	8 249 625	8 252 956	Chrom11	3 099	1 033	117 153.74	5.91
MdGRAS52	MD11G1099100	8 257 519	8 259 770	Chrom11	2 196	732	82 487.76	5.81
MdGRAS53	MD11G1196300	28 120 653	28 122 077	Chrom11	1 422	474	53 013.14	5.90
MdGRAS54	MD11G1204900	29 900 777	29 903 543	Chrom11	1 611	537	59 251.93	5.34
MdGRAS55	MD11G1229600	33 402 652	33 405 823	Chrom11	1 749	583	64 860.12	6.02
MdGRAS56	MD12G1143500	22 266 875	22 270 230	Chrom12	2 253	751	84 128.64	6.77
MdGRAS57	MD12G1176900	25 734 748	25 736 358	Chrom12	1 608	536	60 958.87	4.70
MdGRAS58	MD12G1177000	25 740 719	25 742 332	Chrom12	1 611	537	61 165.83	4.74
MdGRAS59	MD12G1177200	25 763 298	25 764 440	Chrom12	1 140	380	43 170.96	5.48
MdGRAS60	MD12G1196300	27 756 178	27 759 849	Chrom12	2 472	824	88 261.14	6.01
MdGRAS61	MD13G1022100	1 595 594	1 597 541	Chrom13	1 905	635	69 734.67	5.30
MdGRAS62	MD13G1108700	7 807 026	7 808 594	Chrom13	1 566	522	58 256.08	5.67
MdGRAS63	MD13G1155800	12 131 064	12 132 866	Chrom13	1 800	600	67 200.29	4.98
MdGRAS64	MD13G1155900	12 134 055	12 135 380	Chrom13	1 323	441	49 178.14	8.25
MdGRAS65	MD14G1162800	25 718 896	25 720 694	Chrom14	1 641	547	60 843.48	6.05
MdGRAS66	MD15G1180500	14 194 327	14 195 970	Chrom15	1 641	547	59 896.09	5.53
MdGRAS67	MD15G1263000	22 480 624	22 482 399	Chrom15	1 773	591	65 237.96	5.15
MdGRAS68	MD15G1266500	22 905 968	22 907 955	Chrom15	1 644	548	61 187.13	5.69
MdGRAS69	MD15G1294400	27 488 933	27 489 988	Chrom15	1 053	351	39 242.76	5.41
MdGRAS70	MD15G1294600	27 506 241	27 507 587	Chrom15	1 344	448	50 093.24	5.63
MdGRAS71	MD15G1307800	30 733 067	30 734 590	Chrom15	1 521	507	56 852.69	5.38
MdGRAS72	MD15G1353200	42 259 451	42 260 992	Chrom15	1 539	513	56 245.27	5.13
MdGRAS73	MD16G1023300	1 699 006	1 700 925	Chrom16	1 917	639	70 235.28	5.25
MdGRAS74	MD17G1011100	683 081	685 871	Chrom17	2 115	705	78 634.52	5.93
MdGRAS75	MD17G1090800	7 544 673	7 546 001	Chrom17	1 326	442	48 974.67	5.30
MdGRAS76	MD17G1112700	9 646 869	9 650 669	Chrom17	1 653	551	61 949.26	6.39
MdGRAS77	MD17G1128300	11 266 433	11 267 785	Chrom17	1 350	450	48 959.47	5.44
MdGRAS78	MD17G1260700	32 107 773	32 109 527	Chrom17	1 752	584	63 697.47	4.95

基因名 Gene name	基因ID Gene ID	起始位点 Start site	终止位点 End site	染色体 Chromosome	CDS/bp	蛋白质性质Deduce polypetide
基因名 Gene name	基因ID Gene ID	起始位点 Start site	终止位点 End site	染色体 Chromosome	CDS/bp	长度/aa Length	分子量/Da MW	等电点 pI
MdGRAS1	MD00G1043300	7 80 6171	7 809 643	Chrom00	2 424	808	88 359.54	5.80
MdGRAS2	MD00G1112400	23 667 785	23 669 215	Chrom00	1 428	476	53 228.76	5.67
MdGRAS3	MD01G1206800	30 231 215	30 233 158	Chrom01	1 941	647	70 816.40	6.63
MdGRAS4	MD02G1039600	3 216 839	3 218 479	Chrom02	1 638	546	59 826.80	5.63
MdGRAS5	MD02G1148800	12 235 195	12 237 263	Chrom02	1 812	604	66 498.43	5.17
MdGRAS6	MD02G1185900	16 892 351	16 893 874	Chrom02	1 521	507	56 882.60	5.22
MdGRAS7	MD02G1210200	21 832 327	21 834 579	Chrom02	2 250	750	81 783.01	5.80
MdGRAS8	MD02G1295800	35 009 786	35 012 501	Chrom02	1 374	458	51 566.29	5.67
MdGRAS9	MD03G1045700	3 674 587	3 675 963	Chrom03	1 374	458	52 030.80	6.04
MdGRAS10	MD03G1088900	7 330 830	7 332 881	Chrom03	2 049	683	77 700.00	5.52
MdGRAS11	MD03G1089100	7 337 571	7 339 625	Chrom03	2 052	684	78 076.35	6.42
MdGRAS12	MD03G1089200	7 340 949	7 345 605	Chrom03	3 828	1 276	144 938.23	5.93
MdGRAS13	MD03G1089300	7 346 485	7 348 548	Chrom03	2 061	687	77 570.95	5.49
MdGRAS14	MD03G1089700	7 367 711	7 371 218	Chrom03	2 427	809	91 167.55	5.41
MdGRAS15	MD03G1089800	7 380 517	7 382 781	Chrom03	2 208	736	82 619.61	5.28
MdGRAS16	MD03G1169300	22 929 145	22 930 705	Chrom03	738	246	27 338.81	5.90
MdGRAS17	MD03G1188800	25 920 210	25 922 997	Chrom03	1 611	537	59 547.07	5.15
MdGRAS18	MD03G1214700	29 482 170	29 485 609	Chrom03	1 749	583	64 679.63	5.73
MdGRAS19	MD03G1293700	37 023 448	37 025 172	Chrom03	1 722	574	65 184.13	5.94
MdGRAS20	MD04G1046000	5 332 407	5 334 295	Chrom04	1 278	426	47 811.20	5.21
MdGRAS21	MD04G1088100	13 542 153	13 544 592	Chrom04	2 382	794	86 879.60	6.00
MdGRAS22	MD04G1163300	25 473 582	25 475 192	Chrom04	1 608	536	60 735.28	4.67
MdGRAS23	MD04G1163400	25 477 063	25 478 646	Chrom04	1 581	527	59 593.09	4.82
MdGRAS24	MD04G1181500	27 256 030	27 259 385	Chrom04	2 487	829	88 693.53	5.88
MdGRAS25	MD05G1011200	2 343 179	2 345 898	Chrom05	1 737	579	64 386.15	4.96
MdGRAS26	MD05G1040300	6 871 948	6 873 466	Chrom05	1 440	480	54 685.42	6.91
MdGRAS27	MD05G1194300	32 192 340	32 195 288	Chrom05	1 704	568	63 969.95	6.42
MdGRAS28	MD05G1229500	36 294 983	36 296 611	Chrom05	1 626	542	60 679.72	4.86
MdGRAS29	MD05G1360300	47 489 161	47 492 239	Chrom05	1 608	536	60 231.21	5.68
MdGRAS30	MD06G1073500	18 288 623	18 290 677	Chrom06	2 052	684	75 422.74	5.56
MdGRAS31	MD06G1148900	29 171 471	29 173 054	Chrom06	1 581	527	58 725.06	6.37
MdGRAS32	MD07G1030000	2 514 355	2 517 053	Chrom07	1 374	458	51 767.52	5.86
MdGRAS33	MD07G1115800	13 984 309	13 986 546	Chrom07	2 235	745	81 207.10	5.50
MdGRAS34	MD07G1277400	34 273 655	34 275 613	Chrom07	1 956	652	71 075.63	6.27
MdGRAS35	MD08G1167600	19 831 740	19 833 281	Chrom08	1 539	513	56 341.82	5.36
MdGRAS36	MD09G1007200	510 029	513 274	Chrom09	2 112	704	78 249.07	5.82
MdGRAS37	MD09G1103000	7 539 020	7 540 351	Chrom09	1 329	443	49 061.78	5.19
基因名 Gene name	基因ID Gene ID	起始位点 Start site	终止位点 End site	染色体 Chromosome	CDS/bp	蛋白质性质Deduce polypetide
基因名 Gene name	基因ID Gene ID	起始位点 Start site	终止位点 End site	染色体 Chromosome	CDS/bp	长度/aa Length	分子量/Da MW	等电点 pI
MdGRAS38	MD09G1121700	9 383 149	9 386 262	Chrom09	1 659	553	62 323.66	5.74
MdGRAS39	MD09G1137800	10 707 681	10 709 030	Chrom09	1 347	449	48 949.18	5.35
MdGRAS40	MD09G1183000	15 735 881	15 737 539	Chrom09	1 656	552	61 147.14	6.07
MdGRAS41	MD09G1264800	33 799 664	33 801 406	Chrom09	1 740	580	63 166.16	5.09
MdGRAS42	MD10G1011000	1 547 558	1 550 844	Chrom10	1 737	579	64 463.19	5.03
MdGRAS43	MD10G1046300	6 180 700	6 182 115	Chrom10	1 413	471	53 926.17	6.31
MdGRAS44	MD10G1181000	27 411 839	27 413 593	Chrom10	1 542	514	58 030.64	5.91
MdGRAS45	MD10G1336100	41 266 090	41 268 919	Chrom10	1 374	458	51 746.93	6.36
MdGRAS46	MD11G1047900	4 170 134	4 171 525	Chrom11	1 389	463	52 716.63	6.07
MdGRAS47	MD11G1097900	8 118 511	8 120 535	Chrom11	2 022	674	76 651.13	5.45
MdGRAS48	MD11G1098000	8 127 739	8 129 778	Chrom11	2 037	679	77 294.04	5.91
MdGRAS49	MD11G1098100	8 130 580	8 133 055	Chrom11	2 016	672	76 660.73	6.37
MdGRAS50	MD11G1098400	8 139 575	8 141 815	Chrom11	2 166	722	81 747.16	5.92
MdGRAS51	MD11G1099000	8 249 625	8 252 956	Chrom11	3 099	1 033	117 153.74	5.91
MdGRAS52	MD11G1099100	8 257 519	8 259 770	Chrom11	2 196	732	82 487.76	5.81
MdGRAS53	MD11G1196300	28 120 653	28 122 077	Chrom11	1 422	474	53 013.14	5.90
MdGRAS54	MD11G1204900	29 900 777	29 903 543	Chrom11	1 611	537	59 251.93	5.34
MdGRAS55	MD11G1229600	33 402 652	33 405 823	Chrom11	1 749	583	64 860.12	6.02
MdGRAS56	MD12G1143500	22 266 875	22 270 230	Chrom12	2 253	751	84 128.64	6.77
MdGRAS57	MD12G1176900	25 734 748	25 736 358	Chrom12	1 608	536	60 958.87	4.70
MdGRAS58	MD12G1177000	25 740 719	25 742 332	Chrom12	1 611	537	61 165.83	4.74
MdGRAS59	MD12G1177200	25 763 298	25 764 440	Chrom12	1 140	380	43 170.96	5.48
MdGRAS60	MD12G1196300	27 756 178	27 759 849	Chrom12	2 472	824	88 261.14	6.01
MdGRAS61	MD13G1022100	1 595 594	1 597 541	Chrom13	1 905	635	69 734.67	5.30
MdGRAS62	MD13G1108700	7 807 026	7 808 594	Chrom13	1 566	522	58 256.08	5.67
MdGRAS63	MD13G1155800	12 131 064	12 132 866	Chrom13	1 800	600	67 200.29	4.98
MdGRAS64	MD13G1155900	12 134 055	12 135 380	Chrom13	1 323	441	49 178.14	8.25
MdGRAS65	MD14G1162800	25 718 896	25 720 694	Chrom14	1 641	547	60 843.48	6.05
MdGRAS66	MD15G1180500	14 194 327	14 195 970	Chrom15	1 641	547	59 896.09	5.53
MdGRAS67	MD15G1263000	22 480 624	22 482 399	Chrom15	1 773	591	65 237.96	5.15
MdGRAS68	MD15G1266500	22 905 968	22 907 955	Chrom15	1 644	548	61 187.13	5.69
MdGRAS69	MD15G1294400	27 488 933	27 489 988	Chrom15	1 053	351	39 242.76	5.41
MdGRAS70	MD15G1294600	27 506 241	27 507 587	Chrom15	1 344	448	50 093.24	5.63
MdGRAS71	MD15G1307800	30 733 067	30 734 590	Chrom15	1 521	507	56 852.69	5.38
MdGRAS72	MD15G1353200	42 259 451	42 260 992	Chrom15	1 539	513	56 245.27	5.13
MdGRAS73	MD16G1023300	1 699 006	1 700 925	Chrom16	1 917	639	70 235.28	5.25
MdGRAS74	MD17G1011100	683 081	685 871	Chrom17	2 115	705	78 634.52	5.93
MdGRAS75	MD17G1090800	7 544 673	7 546 001	Chrom17	1 326	442	48 974.67	5.30
MdGRAS76	MD17G1112700	9 646 869	9 650 669	Chrom17	1 653	551	61 949.26	6.39
MdGRAS77	MD17G1128300	11 266 433	11 267 785	Chrom17	1 350	450	48 959.47	5.44
MdGRAS78	MD17G1260700	32 107 773	32 109 527	Chrom17	1 752	584	63 697.47	4.95