蓝莓全基因组SSR引物开发及遗传多样性分析

doi:10.13560/j.cnki.biotech.bull.1985.2025-1404

摘要/Abstract

摘要：

目的基于蓝莓品种形态易混淆、缺乏精准鉴定体系的问题，通过全基因组SSR位点的发掘及相关特性分析，开发新的SSR标记引物，为蓝莓种质精准鉴定和遗传多样性评价提供支撑。方法利用Krait软件对‘Draper’基因组SSR位点进行检索分析，并使用Primer 3.0软件设计引物，结合生物信息学技术初步筛选SSR标记引物。逐步扩大蓝莓种质数量进行PCR扩增，采用非变性聚丙烯酰胺凝胶电泳和毛细管电泳相结合的方式，筛选高多态性SSR标记引物，并进行蓝莓种质资源的遗传多样性分析。结果 ‘Draper’基因组SSR位点发掘结果显示，共检测到888 906个SSR位点，以二核苷酸重复类型为主，占总SSR位点数的87.8%，三核苷酸次之，占总SSR位点数的6.34%。经生物信息学初步筛选，获得103对SSR标记引物；经多轮电泳筛选，最终获得17对高多态性SSR标记引物。利用这17对引物对64份蓝莓种质资源进行遗传多样性分析，共检测到204个等位基因，有效等位基因数（Ne）、Shannon’s信息指数（I）平均值分别为6.165和1.956；多态信息量（PIC）平均值为0.985，表明SSR标记多态性较高。UPGMA聚类分析显示，在Nei’s遗传距离0.000 8处，64份蓝莓种质资源可划分为3个聚类群。结论揭示了蓝莓基因组中SSR位点的数量和类型特性。获得17对高多态性SSR引物，可用于区分蓝莓种质资源，且品种间遗传多样性较好。

关键词: 蓝莓, 全基因组, SSR分子标记, SSR引物, 遗传多样性

Abstract:

Objective To address the challenges posed by morphologically ambiguous blueberry cultivars and the lack of a precise identification system, this study aimed to mine genome-wide SSR loci, analyze their characteristics, and develop novel SSR markers, thereby providing a solid foundation for accurate authentication of blueberry germplasm and assessment of genetic diversity. Method SSR loci within the 'Draper' genome were mined and analyzed using Krait software, and primers were designed with Primer 3.0. Preliminary screening of SSR marker primers was performed through bioinformatics analysis. PCR amplification was conducted using a gradually expanded blueberry germplasm collection. A combination of non-denaturing polyacrylamide gel electrophoresis (PAGE) and capillary electrophoresis was employed to screen for highly polymorphic SSR markers. Subsequently, genetic diversity analysis of blueberry germplasm was carried out using the finally selected highly polymorphic primers. Result A total of 888 906 SSR loci were identified in the 'Draper' genome. Dinucleotide repeat motifs were the dominant type, accounting for 87.80% of the total, followed by trinucleotide repeats, which represented 6.34%. Following bioinformatic primer design and preliminary screening, 103 pairs of SSR marker primers were obtained. Subsequent multi-round electrophoretic screening yielded 17 pairs of highly polymorphic SSR markers. These were used to analyze the genetic diversity of 64 blueberry germplasm accessions. A total of 204 alleles were detected. The average effective number of alleles (Ne) was 6.165, the average Shannon’s information index (I) was 1.956, and the average polymorphism information content (PIC) was 0.985, indicating that the developed SSR markers are highly polymorphic. UPGMA cluster analysis showed that the 64 blueberry accessions could be grouped into three distinct clusters at a Nei’s genetic distance of 0.000 8. Conclusion This study characterized the abundance and motif distribution of SSR loci in the blueberry genome. Seventeen highly polymorphic SSR primer pairs were developed, which proved effective for distinguishing blueberry germplasm accessions. Furthermore, the genetic diversity among blueberry cultivars was found to be high.

Key words: Vaccinium spp., whole-genome, SSR molecular marker, SSR primer, genetic diversity

刘思文, 王博伟, 邢金杰, 王静, 李亚东, 孙海悦. 蓝莓全基因组SSR引物开发及遗传多样性分析[J]. 生物技术通报, 2026, 42(11): 134-144.

LIU Si-wen, WANG Bo-wei, XING Jin-jie, WANG Jing, LI Ya-dong, SUN Hai-yue. Genome-wide Development of SSR Primers and Genetic Diversity Analysis in Blueberry[J]. Biotechnology Bulletin, 2026, 42(11): 134-144.

图/表 11

参考文献 40

[1]	Duan YM, Tarafdar A, Chaurasia D, et al. Blueberry fruit valorization and valuable constituents: a review [J]. Int J Food Microbiol, 2022, 381: 109890.
[2]	Jin ZW, Huang HZ, Huang HH, et al. The delayed senescence in harvested blueberry by hydrogen-based irrigation is functionally linked to metabolic reprogramming and antioxidant machinery [J]. Food Chem, 2024, 453: 139563.
[3]	Yu YY, Xu JD, Huang TX, et al. Combination of beneficial bacteria improves blueberry production and soil quality [J]. Food Sci Nutr, 2020, 8(11): 5776-5784.
[4]	李亚东, 刘成, 魏鑫, 等. 2024年中国蓝莓产业发展报告 [J]. 吉林农业大学学报, 2025, 47(1): 1-14.
	Li YD, Liu C, Wei X, et al. Development report of 2024 China blueberry industry [J]. J Jilin Agric Univ, 2025, 47(1): 1-14.
[5]	雷艳珍. 杂交种授权亲本品种权保护问题研究‒‒兼评新《植物新品种保护条例》第7条第3款第3项 [J]. 法治研究, 2025(6): 131-142.
	Lei YZ. Research on the protection of parental variety rights for hybrid varieties [J]. Res Rule Law, 2025(6): 131-142.
[6]	Cipriani G, Marrazzo M, Di Gaspero G, et al. A set of microsatellite markers with long core repeat optimized for grape (Vitis spp.) genotyping [J]. BMC Plant Biol, 2008, 8(1): 127.
[7]	Daudi H, Shimelis H, Mathew I, et al. Genetic diversity and population structure of groundnut (Arachis hypogaea L.) accessions using phenotypic traits and SSR markers: implications for rust resistance breeding [J]. Genet Resour Crop Evol, 2021, 68(2): 581-604.
[8]	Baruca Arbeiter A, Hladnik M, Jakše J, et al. First set of microsatellite markers for immortelle (Helichrysum italicum (Roth) G. Don): a step towards the selection of the most promising genotypes for cultivation [J]. Ind Crops Prod, 2021, 162: 113298.
[9]	Boutsika A, Sarrou E, Cook CM, et al. Evaluation of parsley (Petroselinum crispum) germplasm diversity from the Greek Gene Bank using morphological, molecular and metabolic markers [J]. Ind Crops Prod, 2021, 170: 113767.
[10]	Savadi S, Muralidhara BM, Venkataravanappa V, et al. De novo transcriptome assembly and its utility in development and characterization of the first set of genic SSR markers in cashew [J]. Ind Crops Prod, 2022, 189: 115734.
[11]	岳林清, 吴怡超, 何星星, 等. 李资源果实主要性状的SSR标记关联分析及指纹图谱构建 [J]. 园艺学报, 2024, 51(11): 2495-2509.
	Yue LQ, Wu YC, He XX, et al. SSR marker association analysis and fingerprint construction of main traits of plum fruit [J]. Acta Hortic Sin, 2024, 51(11): 2495-2509.
[12]	牛锐敏, 黄小晶, 沈甜, 等. 基于SSR标记的葡萄种质指纹图谱构建和遗传多样性分析 [J]. 种子, 2025, 44(4): 106-114.
	Niu RM, Huang XJ, Shen T, et al. Fingerprints construction and genetic diversity analysis of Vitis vinifera germplasm resources based on SSR markers [J]. Seed, 2025, 44(4): 106-114.
[13]	闫锦源, 郑斌, 王松标, 等. 基于荧光SSR标记的杧果控制授粉实生后代父本鉴定 [J]. 果树学报, 2025, 42(6): 1190-1199.
	Yan JY, Zheng B, Wang SB, et al. Paternal identification of mango controlled-pollination seedling proge-nies based on fluorescent SSR markers [J]. J Fruit Sci, 2025, 42(6): 1190-1199.
[14]	董清华, 王西成, 赵密珍, 等. 草莓EST-SSR标记开发及在品种遗传多样性分析中的应用 [J]. 中国农业科学, 2011, 44(17): 3603-3612.
	Dong QH, Wang XC, Zhao MZ, et al. Development of EST-derived SSR markers and their application in strawberry genetic diversity analysis [J]. Sci Agric Sin, 2011, 44(17): 3603-3612.
[15]	王静, 赵密珍, 于红梅, 等. 草莓EST-SSR标记的开发与应用 [J]. 果树学报, 2011, 28(4): 721-726.
	Wang J, Zhao MZ, Yu HM, et al. Development and utilization of EST-SSR marker in strawberry [J]. J Fruit Sci, 2011, 28(4): 721-726.
[16]	Song YP, Jiang XB, Zhang M, et al. Differences of EST-SSR and genomic-SSR markers in assessing genetic diversity in poplar [J]. For Stud China, 2012, 14(1): 106.
[17]	苗立祥, 杨肖芳, 张豫超, 等. 基于草莓全基因组SSR标记的开发和应用 [J]. 分子植物育种, 2021, 19(4): 1210-1222.
	Miao LX, Yang XF, Zhang YC, et al. Development of genome-wide SSR marker and application in strawberry [J]. Mol Plant Breed, 2021, 19(4): 1210-1222.
[18]	刘嘉艺, 岳俊阳, 刘永胜. 基于毛花猕猴桃基因组的性别相关SSR分子标记的开发 [J]. 合肥工业大学学报: 自然科学版, 2022, 45(8): 1135-1138, 1146.
	Liu JY, Yue JY, Liu YS. Development of sex-related SSR molecular markers based on the genome of Actinidia eriantha [J]. J Hefei Univ Technol Nat Sci, 2022, 45(8): 1135-1138, 1146.
[19]	Liu YC, Liu S, Liu DC, et al. Exploiting EST databases for the development and characterization of EST-SSR markers in blueberry (Vaccinium) and their cross-species transferability in Vaccinium spp [J]. Sci Hortic, 2014, 176: 319-329.
[20]	王利平, 沈颖, 陈荣锋, 等. 蓝莓基因组SSR分子标记开发 [J]. 浙江农业科学, 2023, 64(6): 1503-1506.
	Wang LP, Shen Y, Chen RF, et al. Development of SSR molecular markers of blueberry genome [J]. J Zhejiang Agric Sci, 2023, 64(6): 1503-1506.
[21]	Zeng T, He ZJ, He JF, et al. The telomere-to-telomere gap-free reference genome of wild blueberry (Vaccinium duclouxii) provides its high soluble sugar and anthocyanin accumulation [J]. Hortic Res, 2023, 10(11): uhad209.
[22]	Wang ZJ, Zhang WC, Zhou YY, et al. Genetic and epigenetic signatures for improved breeding of cultivated blueberry [J]. Hortic Res, 2024, 11(7): uhae138.
[23]	Gill K, Negi S, Kumar P, et al. Improved genomic DNA extraction from citrus species using a modified CTAB method[J]. Mol Biol Rep, 2025, 52(1): 638.
[24]	Du LM, Zhang C, Liu Q, et al. Krait: An ultrafast tool for genome-wide survey of microsatellites and primer design [J]. Bioinformatics, 2018, 34(4): 681-683.
[25]	樊文强, 盖红梅, 孙鑫, 等. SSR数据格式转换软件DataFormater [J]. 分子植物育种, 2016, 14(1): 265-270.
	Fan WQ, Gai HM, Sun X, et al. DataFormater, a software for SSR data formatting to develop population genetics analysis [J]. Mol Plant Breed, 2016, 14(1): 265-270.
[26]	Zhao MQ, Shu GP, Hu YH, et al. Pattern and variation in simple sequence repeat (SSR) at different genomic regions and its implications to maize evolution and breeding [J]. BMC Genom, 2023, 24: 136.
[27]	孙海悦, 郭瑞雪, 徐鉴, 等. 越橘属植物EST-SSR分子标记开发及遗传多样性分析 [J]. 植物生理学报, 2018, 54(3): 429-440.
	Sun HY, Guo RX, Xu J, et al. Development of EST-SSR molecular markers for Vaccinium and genetic diversity analysis [J]. Plant Physiol J, 2018, 54(3): 429-440.
[28]	范佳妮, 蒋丽娟, 李培旺, 等. 无患子全基因组SSR标记开发及遗传多样性分析[J]. 分子植物育种, 2025. .
	Fan JN, Jiang LJ, Li PW, et al. Development of SSR markers and genetic diversity analysis of Sapindus saponaria genome [J]. Mol Plant Breed, 2025. .
[29]	朱跃, 张敏, 张成, 等. 桂花全基因组SSR特征分析与引物开发 [J]. 南京林业大学学报: 自然科学版, 2025, 49(5): 174-182.
	Zhu Y, Zhang M, Zhang C, et al. Genome wide SSR analysis and primer development of Osmanthus fragrans [J]. J Nanjing For Univ Nat Sci Ed, 2025, 49(5): 174-182.
[30]	张咏琦, 王超, 许林林, 等. 野生二倍体草莓的SSR分子标记开发及核心种质的构建 [J]. 园艺学报, 2023, 50(11): 2365-2375.
	Zhang YQ, Wang C, Xu LL, et al. Development of SSR molecular markers and construction of core collection of wild diploid strawberry [J]. Acta Hortic Sin, 2023, 50(11): 2365-2375.
[31]	Zhang CC, Lin P, Yao XH, et al. Genome-wide development of genomic SSR markers, polymorphism analysis, and dosage-aware SSR fingerprinting in hexaploid Camellia Oleifera [J]. BMC Plant Biol, 2025, 25: 1627.
[32]	蒋爽, 张学英, 安海山, 等. 枇杷全基因组SSR标记开发及其多态性研究 [J]. 园艺学报, 2021, 48(5): 1013-1022.
	Jiang S, Zhang XY, An HS, et al. Development and analysis of polymorphism of SSR markers in the whole genome of loquat [J]. Acta Hortic Sin, 2021, 48(5): 1013-1022.
[33]	郭瑞雪. 越橘EST-SSR分子标记的开发及遗传多样性分析 [D]. 长春: 吉林农业大学, 2014.
	Guo RX. Development of EST-derived SSR markers and genetic diversity analysis in Vaccinium [D]. Changchun: Jilin Agricultural University, 2014.
[34]	方茜, 张园园, 杨钰婷, 等. 越橘叶片转录组SSR发掘及其多态性研究 [J]. 园艺学报, 2018, 45(7): 1359-1370.
	Fang Q, Zhang YY, Yang YT, et al. SSR mining and polymorphism analysis in leaf transcriptome of blueberry [J]. Acta Hortic Sin, 2018, 45(7): 1359-1370.
[35]	俞新华, 李超, 哈雪姣, 等. 西瓜品种SSR指纹库构建及应用 [J]. 蔬菜, 2025(12): 37-46.
	Yu XH, Li C, Ha XJ, et al. Construction and application of SSR fingerprint database for watermelon varieties [J]. Vegetables, 2025(12): 37-46.
[36]	吴仕蔓, 娄兵海, 陈传武, 等. 应用SSR荧光标记法构建22个柚类品种的分子身份证 [J]. 果树学报, 2023, 40(4): 605-614.
	Wu SM, Lou BH, Chen CW, et al. Establishment of molecular identity of 22 pomelo varieties using fluores-cent labeled SSR markers [J]. J Fruit Sci, 2023, 40(4): 605-614.
[37]	崔建民, 刘红霞, 邹荣仟, 等. 越橘种质资源遗传多样性和亲缘关系研究 [J]. 果树学报, 2010, 27(3): 373-378.
	Cui JM, Liu HX, Zou RQ, et al. Study on genetic diversity and phylogenetic relationship of germplasm resources about Vaccinium spp [J]. J Fruit Sci, 2010, 27(3): 373-378.
[38]	郭照东, 夏秀英, 安利佳, 等. 基于SSR标记的越橘亲缘关系分析及品种鉴定 [J]. 植物遗传资源学报, 2015, 16(5): 1020-1026.
	Guo ZD, Xia XY, An LJ, et al. Genetic relationship analysis and cultivar identification of blueberry based on SSR markers [J]. J Plant Genet Resour, 2015, 16(5): 1020-1026.
[39]	Haque MS, Saha NR, Islam MT, et al. Screening for drought tolerance in wheat genotypes by morphological and SSR markers [J]. J Crop Sci Biotechnol, 2021, 24(1): 27-39.
[40]	Tailor S, Bykova NV, Igamberdiev AU, et al. Structural pattern and genetic diversity in blueberry (Vaccinium) clones and cultivars using EST-PCR and microsatellite markers [J]. Genet Resour Crop Evol, 2017, 64(8): 2071-2082.

编号 Code	品种名 Cultivar name	编号 Code	品种名 Cultivar name	编号 Code	品种名 Cultivar name	编号 Code	品种名 Cultivar name
1	Draper	17	蓝塔	33	努伊	49	Emil
2	利宝	18	泽西	34	伯灵顿	50	L25
3	甜心	19	雷格西	35	艾玛	51	1502
4	Biloxi	20	博尼法西	36	AF1	52	H5
5	X13	21	赫伯特	37	AF2	53	Y42
6	北春	22	布里吉塔	38	AF3	54	F6
7	North sky	23	托柔	39	Small fruit	55	L29
8	North blue	24	日升	40	Jgdrzej	56	122
9	北陆	25	奥林匹克	41	HL2	57	L11
10	慧蓝	26	早蓝	42	HL3	58	49
11	圣云	27	都克	43	HL4	59	O2
12	蓝金	28	晚蓝	44	HL5	60	L45
13	普特	29	艾玛蓝	45	HL9	61	YOU
14	蓝丰	30	瑞卡	46	贝尔	62	DS01
15	H7	31	考林	47	格瑞萨	63	DS04
16	齐伯瓦	32	Record	48	北卫	64	D15

编号 Code	品种名 Cultivar name	编号 Code	品种名 Cultivar name	编号 Code	品种名 Cultivar name	编号 Code	品种名 Cultivar name
1	Draper	17	蓝塔	33	努伊	49	Emil
2	利宝	18	泽西	34	伯灵顿	50	L25
3	甜心	19	雷格西	35	艾玛	51	1502
4	Biloxi	20	博尼法西	36	AF1	52	H5
5	X13	21	赫伯特	37	AF2	53	Y42
6	北春	22	布里吉塔	38	AF3	54	F6
7	North sky	23	托柔	39	Small fruit	55	L29
8	North blue	24	日升	40	Jgdrzej	56	122
9	北陆	25	奥林匹克	41	HL2	57	L11
10	慧蓝	26	早蓝	42	HL3	58	49
11	圣云	27	都克	43	HL4	59	O2
12	蓝金	28	晚蓝	44	HL5	60	L45
13	普特	29	艾玛蓝	45	HL9	61	YOU
14	蓝丰	30	瑞卡	46	贝尔	62	DS01
15	H7	31	考林	47	格瑞萨	63	DS04
16	齐伯瓦	32	Record	48	北卫	64	D15

名称 Name	正向引物序列 Forward primer sequences （5′‒3′）	反向引物序列 Reverse primer sequence （5′‒3′）	退火温度 Tm （℃）	预期产物 Expected size （bp）	荧光类型 Fluorescent types
spSSR03	CTGTGTCGACCTCTCCAACG	TGAGATCGGTGGCTTTCTCG	57	242	FAM
spSSR04	CTTGGCGATCTCCTCCTTCC	TCAAGGAACAACAGGCCTCC	55	320	HEX
spSSR05	GCTTCCGGTACCTCTCTTCG	TCATCCCGCACAAGAGTTCC	57	283	HEX
spSSR08	ATCTGAAGTCTCCAACGGCG	GCTAACCCGAGTCTGGTTCC	55	262	HEX
spSSR09	TGTGGCTGGAGATGTTGACG	CAAAATGCGATTCCTCCTCCC	57	236	FAM
spSSR11	AAAGGGGACAAGCAACTCCC	CACCACTTCCGGGATAGTGC	57	252	FAM
spSSR12	ACGACGTGAGAAACCACTCC	CTATCCCTCAACCGCGATGG	57	262	HEX
spSSR13	GAGAGAGAGCCAGTCCTTGC	ACGACGAACCAGACGAAACC	57	194	FAM
spSSR14	ACGACGAACCAGACGAAACC	GAGAGAGAGCCAGTCCTTGC	59	209	FAM
spSSR15	TGAACCAGACGAAACCCACC	GAGAGAGAGCCAGTCCTTGC	59	178	FAM
spSSR16	CAAGTCGACAACATGCTCCG	TTTACAAGCACATACGCGCG	59	218	FAM
spSSR18	CTTGGCGATCTCCTCCTTCC	TCAAGGAACAACAGGCCTCC	59	332	HEX
spSSR19	TTCCAAAACCACCAAACGCC	AGAGCAATGTGGTAGCCAGG	55	326	HEX
spSSR20	GGCTTACAGCTCCTCCATGG	GGTGATGAGAGAGAGCCAGG	59	175	FAM
spSSR21	CTATCCCTCAACCGCGATGG	ACGACGTGAGAAACCACTCC	59	268	HEX
spSSR22	TGTGCATCCAAACCTTGAGC	TGTGGTGGTGGTTATGGAGG	59	238	FAM
spSSR23	GAGGCGGTAGTACATGTGGG	CTCTCAACTCCCCTTCAGCC	57	340	HEX

名称 Name	正向引物序列 Forward primer sequences （5′‒3′）	反向引物序列 Reverse primer sequence （5′‒3′）	退火温度 Tm （℃）	预期产物 Expected size （bp）	荧光类型 Fluorescent types
spSSR03	CTGTGTCGACCTCTCCAACG	TGAGATCGGTGGCTTTCTCG	57	242	FAM
spSSR04	CTTGGCGATCTCCTCCTTCC	TCAAGGAACAACAGGCCTCC	55	320	HEX
spSSR05	GCTTCCGGTACCTCTCTTCG	TCATCCCGCACAAGAGTTCC	57	283	HEX
spSSR08	ATCTGAAGTCTCCAACGGCG	GCTAACCCGAGTCTGGTTCC	55	262	HEX
spSSR09	TGTGGCTGGAGATGTTGACG	CAAAATGCGATTCCTCCTCCC	57	236	FAM
spSSR11	AAAGGGGACAAGCAACTCCC	CACCACTTCCGGGATAGTGC	57	252	FAM
spSSR12	ACGACGTGAGAAACCACTCC	CTATCCCTCAACCGCGATGG	57	262	HEX
spSSR13	GAGAGAGAGCCAGTCCTTGC	ACGACGAACCAGACGAAACC	57	194	FAM
spSSR14	ACGACGAACCAGACGAAACC	GAGAGAGAGCCAGTCCTTGC	59	209	FAM
spSSR15	TGAACCAGACGAAACCCACC	GAGAGAGAGCCAGTCCTTGC	59	178	FAM
spSSR16	CAAGTCGACAACATGCTCCG	TTTACAAGCACATACGCGCG	59	218	FAM
spSSR18	CTTGGCGATCTCCTCCTTCC	TCAAGGAACAACAGGCCTCC	59	332	HEX
spSSR19	TTCCAAAACCACCAAACGCC	AGAGCAATGTGGTAGCCAGG	55	326	HEX
spSSR20	GGCTTACAGCTCCTCCATGG	GGTGATGAGAGAGAGCCAGG	59	175	FAM
spSSR21	CTATCCCTCAACCGCGATGG	ACGACGTGAGAAACCACTCC	59	268	HEX
spSSR22	TGTGCATCCAAACCTTGAGC	TGTGGTGGTGGTTATGGAGG	59	238	FAM
spSSR23	GAGGCGGTAGTACATGTGGG	CTCTCAACTCCCCTTCAGCC	57	340	HEX

名称 Name	等位基因数Na	有效等位基因数Ne	Shannon’s信息指数I	观测杂合度Ho	期望杂合度He	无偏期望杂合度uHe	多态性信息量PIC
spSSR03	6	3.626	1.405	0.714	0.724	0.730	0.983
spSSR04	16	10.431	2.520	0.750	0.904	0.911	0.990
spSSR05	5	2.602	1.083	0.500	0.616	0.623	0.978
spSSR08	9	4.958	1.786	0.583	0.798	0.807	0.986
spSSR09	9	3.292	1.475	1.000	0.696	0.702	0.973
spSSR11	10	4.698	1.789	1.000	0.787	0.794	0.982
spSSR12	9	4.898	1.735	1.000	0.796	0.803	0.984
spSSR13	17	5.737	2.154	1.000	0.826	0.832	0.988
spSSR14	15	7.008	2.188	1.000	0.857	0.864	0.990
spSSR15	19	10.139	2.558	0.800	0.901	0.908	0.990
spSSR16	14	9.806	2.383	0.667	0.898	0.905	0.989
spSSR18	16	10.711	2.542	0.200	0.907	0.914	0.990
spSSR19	13	6.122	2.099	1.000	0.837	0.844	0.985
spSSR20	13	5.306	1.993	1.000	0.812	0.818	0.980
spSSR21	11	3.924	1.631	1.000	0.745	0.751	0.981
spSSR22	10	5.760	1.913	1.000	0.826	0.833	0.983
spSSR23	12	5.796	1.992	1.000	0.827	0.834	0.983
平均Mean	12	6.165	1.956	0.836	0.809	0.816	0.985