基于CAPS标记的甜瓜单果重相关性状QTL分析

doi:10.3864/j.issn.0578-1752.2019.09.011

摘要/Abstract

摘要：

【目的】基于甜瓜全基因组重测序技术,挖掘SNP位点并开发CAPS标记,构建遗传连锁图谱。初步为甜瓜单果重相关性状进行QTL定位,为甜瓜单果重相关基因挖掘奠定理论基础。【方法】选取栽培甜瓜品系M4-130为母本,野生甜瓜品系X207为父本,构建F_2:3群体材料。对甜瓜果实单果重、果实长宽、果肉厚度、果形指数及可溶性固形物含量等性状进行相关性分析。对亲本材料进行20×深度重测序,利用BWA、SAMTools、VCFTools等软件在全基因组范围内挖掘双亲SNP位点,利用SNP2CAPS软件结合甜瓜基因组上的限制性内切酶酶切位点开发CAPS标记,建立遗传连锁图谱。最后采用复合区间作图法对甜瓜果实单果重、果实长宽、果肉厚度、果形指数及可溶性固形物含量进行QTL分析。【结果】单果重与果实长度、果实宽度及果肉厚度呈显著相关。开发得到可利用CAPS标记185个,构建一张包含12个连锁群的遗传连锁图谱,覆盖总长度为1 600.45 cM,标记间的平均遗传距离8.65 cM。定位发现与单果重相关QTL位点7个（FW3.1、FW4.1、FW5.1、FW6.1、FW8.1、FW8.2、FW11.1）,与果实长度相关QTL位点7个（FL2.1、FL3.1、FL4.1、FL5.1、FL6.1、FL8.1、FL11.1）,与果实宽度相关QTL位点5个（FWID3.1、FWID4.1、FWID8.1、FWID10.1、FWID11.1）,与果肉厚度相关QTL位点2个（FT6.1、FT11.1）,与果形指数相关QTL位点1个（FS2.1）,与可溶性固形含量相关QTL位点2个（SS6.1、SS12.1）。【结论】获得了24个QTL位点。确定了一个主效QTL位点FW8.1,贡献率高达25.8774%,LOD值为16.8746。发现单果重与果实长度、果实宽度及果肉厚度密切相关,定位区间相同或相邻,紧密集中在3、4、5、6、8、11染色体上。

关键词: 甜瓜, 单果重, 遗传连锁图谱, QTL

Abstract:

【Objective】 In our study, based on melon genome-wide resequencing technology, SNP loci were explored and CAPS markers were developed to construct a genetic linkage map. The preliminary QTL analysis of traits related to single fruit weight in melon was carried out, which laid a theoretical foundation for the detection of genes related to single fruit weight traits in melon.【Method】F_2:3 population materials were constructed from X207 and M4-130. The cultivated melon accession M4-130 was selected as female parent and wild melon accession X207 as male parent. Correlation analysis was carried out for fruit traits, including single fruit weight, fruit length, fruit width, pulp thickness, fruit shape index and soluble solids content in melon fruit. Parents were resequenced at 20× depth. SNP loci between parents were extracted in the whole genome by the use of BWA, SAMTools, VCFTools and other software. CAPS markers were developed by using SNP2CAPS software and combined with restriction endonuclease digestion sites in melon genome for genetic linkage map construction. Finally, QTLs of single fruit weight, fruit length, fruit width, pulp thickness, fruit shape index and soluble solids content in melon fruit were analyzed by composite interval mapping.【Result】In this experiment, single fruit weight significantly correlated with fruit length, fruit width and pulp thickness. 185 CAPS markers were developed and subsequently used for genetic linkage map construction. The genetic linkage map contained 12 linkage groups covering a total length of 1 600.45 cM and average distance of 8.65 cM. 7 QTLs (FW3.1, FW4.1, FW5.1, FW6.1, FW8.1, FW8.2 and FW11.1) were related to the single fruit weight. 7 QTLs (FL2.1, FL3.1, FL4.1, FL5.1, FL6.1, FL8.1 and FL11.1) were related to fruit length. 5 QTLs (FWID3.1, FWID4.1, FWID8.1, FWID10.1 and FWID11.1) were related to fruit width. 2 QTLs (FT6.1 and FT11.1) were related to pulp thickness. 1 QTLs (FS2.1) was related to fruit shape index. 2 QTLs (SS6.1 and SS12.1) were related to soluble solids content.【Conclusion】Twenty-three QTLs in total were identified. A major QTL FW8.1 was identified with phenotypic variance contribution rate of 25.8774% and the LOD value was 16.8746. It was also found that single fruit weight was closely related to fruit length, fruit width and pulp thickness. Location intervals were same or adjacent, closely concentrated on chromosomes 3, 4, 5, 6, 8 and 11.

Key words: melon, fruit weight, genetic linkage map, QTL

刘相玉, 张裕舒, 刘柳, 刘识, 高鹏, 王迪, 王学征. 基于CAPS标记的甜瓜单果重相关性状QTL分析[J]. 中国农业科学, 2019, 52(9): 1601-1613.

LIU XiangYu, ZHANG YuShu, LIU Liu, LIU Shi, GAO Peng, WANG Di, WANG XueZheng. The QTL Analysis of Single Fruit Weight Associated Traits in Melon Based on CAPS Markers[J]. Scientia Agricultura Sinica, 2019, 52(9): 1601-1613.

0
/ / 推荐

导出引用管理器 EndNote|Reference Manager|ProCite|BibTeX|RefWorks

链接本文: https://www.chinaagrisci.com/CN/10.3864/j.issn.0578-1752.2019.09.011

https://www.chinaagrisci.com/CN/Y2019/V52/I9/1601

图/表 12

图1

表1

图2

图3

表2

图4

图5

表3

图6

表4

甜瓜果实相关性状QTL分析及其遗传效力"

性状 Trait	位点 QTL	相邻标记 Adjacent markers	图谱位置 Position	标记间距 Marker range	LOD	贡献率（%） R²	加性效应 Additive	显性效应 Dominance
单果重 Fruit weight	FW3.1	3-6670—3-0640	66	65.23—74.69	3.0173	2.7054	39.2379	32.2348
	FW4.1	4-9382—4-1311	31	27.25—31.68	3.2739	12.3435	39.6980	-14.1269
	FW5.1	5-9232—5-5773	108	105.17—108.46	2.9070	9.3215	-31.2002	23.5592
	FW6.1	6-1444—6-6615	160	106.62—160.37	3.9121	14.4755	40.7147	-20.9489
	FW8.1	8-2883—8-5972	101	98.60—104.93	16.8746	25.8774	-1.2134	184.5905
	FW8.2	8-0306—8-2169	71	65.29—72.09	3.3297	3.6869	3.6869	9.8677
	FW11.1	11-3692—11-1432	48	41.93—56.06	4.6085	18.8628	48.3023	-18.9123
果实长度 Fruit length	FL2.1	2-2347—2-1803	104	100.56—109.19	6.7436	17.0180	-1.3111	0.2646
	FL3.1	3-6670—3-0640	66	65.23—74.69	3.0160	15.1015	1.1146	0.9123
	FL4.1	4-9382—4-1311	30	27.25—31.68	4.2371	9.9603	0.9998	-0.2666
	FL5.1	5-9232—5-5773	107	105.17—108.46	2.6505	5.1442	-0.6258	0.5634
	FL6.1	6-1444—6-6615	148	106.62—160.37	3.8379	14.1998	1.1898	-0.0833
	FL8.1	8-8589—8-8413	14	00.00—14.22	2.8085	19.9334	1.3147	-0.5710
	FL11.1	11-8315—11-3692	35	30.15—41.93	6.1904	15.7421	1.2632	-0.0376
果实宽度 Fruit width	FWID3.1	3-1192—3-0090	34	18.50—46.50	3.0311	11.3927	0.6226	0.0665
	FWID4.1	4-1311—4-2962	34	31.68—42.76	3.2242	10.1598	0.7384	0.1435
	FWID8.1	8-0306—8-2169	71	65.29—72.09	5.3190	18.4669	0.7200	-0.3225
	FWID10.1	10-3838—10-0417	1	0.00—9.81	4.0028	13.2077	0.6771	0.0365
	FWID11.1	11-8315—11-3692	38	30.15—41.93	4.5617	15.8568	0.9479	-0.0865
果肉厚度 Fruit thickness	FT6.1	6-1444—6-6615	151	106.62—160.37	2.8921	12.0802	0.1414	-0.2006
果肉厚度 Fruit thickness	FT11.1	11-8315—11-3692	35	30.15—41.93	4.3730	15.2309	0.2292	-0.0031
果型指数 Fruit shape index	FS2.1	2-1803—2-2808	112	109.19—112.55	4.6355	22.0413	-0.1131	0.0923
可溶性固形物 Soluble solids	SS6.1	6-1444—6-6615	146	106.62—160.37	2.9412	16.4299	1.2740	0.2516
可溶性固形物 Soluble solids	SS12.1	12-5356—12-4957	9	0.00—19.61	3.3407	10.0947	0.1159	-1.4154

表4

表5

223个基因功能注释"

GO一级类别 GO Level I Category	GO数据库二级分类编号 GO Level II Category	GO分类注释 Annotation of II level GO term	基因数量 Gene number
生物学过程 Biological process	GO: 0008152	代谢过程 Metabolic process	91
	GO: 0009987	细胞过程 Cellular process	83
	GO: 0044699	单体进程 Single-organism process	65
	GO: 0065007	生物调控 Biological regulation	31
	GO: 0050896	应激反应 Response to stimulus	20
	GO: 0071840	细胞成分组织与合成 Cellular component organization or biogenesis	19
	GO: 0051179	定位 Localization	16
	GO: 0032502	发展过程 Developmental process	11
	GO: 0032501	多细胞生物过程 Multicellular organismal process	9
	GO: 0000003	生殖过程 Reproduction	6
	GO: 0022414	再生过程 Reproductive process	6
	GO: 0023052	信号 Signaling	4
	GO: 0040007	发育过程 Growth	3
	GO: 0051704	多生物过程 Multi-organism process	3
	GO: 0022610	生物粘附 Biological adhesion	1
	GO: 0098754	去毒过程 Detoxification	1
分子功能 Molecular function	GO: 0005488	联结 Binding	95
	GO: 0003824	催化活性 Catalytic activity	81
	GO: 0005215	转运活性 Transporter activity	8
	GO: 0001071	核酸结合转录因子活性 Nucleic acid binding transcription factor activity	4
	GO: 0098772	分子功能调控 Molecular function regulator	4
	GO: 0005198	结构分子活性 Structural molecule activity	3
	GO: 0016209	抗氧化活性 Antioxidant activity	3
	GO: 0000988	蛋白质结合转录活性 Transcription factor activity, protein binding	1
	GO: 0004871	信号传感器活性 Signal transducer activity	1
	GO: 0060089	分子转导活性 Molecular transducer activity	1
细胞组分（Cellular Component）	GO: 0005623	细胞 Cell	73
	GO: 0044464	细胞组分 Cell part	73
	GO: 0043226	细胞器 Organelle	49
	GO: 0016020	细胞膜 Membrane	30
	GO: 0032991	大分子复合物 Macromolecular complex	26
	GO: 0044425	细胞膜组分 Membrane part	20
	GO: 0044422	细胞器组分 Organelle part	15
	GO: 0005576	细胞外区 Extracellular region	4
	GO: 0030054	细胞连接 Cell junction	4
	GO: 0055044	共质体 Symplast	4
	GO: 0031974	膜封闭的管腔 Membrane-enclosed lumen	3
	GO: 0019012	病毒 Virion	2
	GO: 0044423	病毒组分 Virion part	2

表5

表6

参考文献 33

[1]	VERZERA A, DIMA G, TRIPODI G, CONDURSO C, CRINO P, ROMANO D, MAZZAGLIA A, LANZA C M, RESTUCCIA C, PARATORE A . Aroma and sensory quality of honeydew melon fruits (Cucumis melo L. subsp. melo var. inodorus H. Jacq.) in relation to different rootstocks. Scientia Horticulturae, 2014,169(11):118-124.
[2]	DANTAS D D C, MEDIROS J F, FREIRE A G . Produção e qualidade do meloeiro cultivado com filmes plásticos em respostas à lâmina de irrigação. Revista Ciência Agronômica, 2011,42(3):652-661. doi: 10.1590/S1806-66902011000300011
[3]	董玉梅, 王崇启, 赵西, 肖守华, 陈伟, 孙建磊, 焦自高 . 不同嫁接方式对厚皮甜瓜植株生长及果实性状的影响. 中国瓜菜, 2017,30(9):34-37.
	DONG Y M, WANG C Q, ZHAO X, XIAO S H, CHEN W, SUN J L, JIAO Z G . Influences of different grafting methods on the plant growth and fruit characters of muskmelon. Zhongguo Gua-cai, 2017,30(9):34-37. (in Chinese)
[4]	于翠香, 韩忠才, 王占海, 张海燕, 杨贵春 . 甜瓜果实性状遗传规律的研究进展. 东北农业科学, 2016,41(3):91-94.
	YU C X, HAN Z C, WANG Z H, ZHANG H Y, YANG G C . Advance of studies on genetic law of melon fruit characters. Journal of Northeast Agricultural Sciences, 2016,41(3):91-94. (in Chinese)
[5]	陈克农, 张红卓, 袁丽伟, 盛云燕 . 甜瓜果形指数的F₂代遗传研究. 北方园艺, 2016(19):1-2.
	CHEN K N, ZHANG H Z, YUAN L W, SHENG Y Y . Genetic inheritance analysis on melon index in F₂ generation. Northern Horticulture, 2016(19):1-2. (in Chinese)
[6]	张宁 . 甜瓜远缘群体果实主要性状遗传分析及遗传图谱构建[D]. 杨凌: 西北农林科技大学, 2014.
	ZHANG N . Genetic analysis of main fruit traits and construction of the genetic linkage map for interspecific population of melon[D]. Yangling: Northwest A&F University, 2014. ( in Chinese)
[7]	BAUDRACCO-ARNAS S, PITRAT M . A genetic map of melon (Cucumis melo L.) with RFLP, RAPD, isozyme, disease resistance and morphological markers. Theoretical and Applied Genetics, 1996,93(1/2):57-64.
[8]	PÉRIN C, HAGEN L, De CONTO V, KATZIR N, DANIN-POLEG Y, PORTNOY V, BAUDRACCO-ARNAS S, CHADOEUF J, DOGIMONT C, PITRAT M . A reference map of Cucumis melo based on two recombinant inbred line populations. Theoretical and Applied Genetics, 2002,104(6):1017-1034.
[9]	CUEVAS H E, STAUB J E, SIMON P W, ZALAPA J E . A consensus linkage map identifies genomic regions controlling fruit maturity and beta-carotene associated flesh color in melon (Cucumis melo L.). Theoretical and Applied Genetics, 2009,119(4):741-756.
[10]	FUKINO N, OHARA T, SUGIYAMA M, KUBO N, HIRAI M, SAKATA Y, MATSUMOTO S . Mapping of a gene that confers short lateral branching ( slb) in melon(Cucumis melo L.). Euphytica, 2012,187:133-143.
[11]	ZALAPA J E, STAUB J E, MCCREIGHT J D, CHUNG S M, CUEVAS H . Detection of QTL for yield-related traits using recombinant inbred lines derived from exotic and elite US Western Shipping melon germplasm. Theoretical & Applied Genetics, 2007,114(7):1185-1201.
[12]	RAMAMURTHY R K, WATERS B M . Identification of fruit quality and morphology QTLs in melon (Cucumis melo) using a population derived from flexuosus and cantalupensis botanical groups. Euphytica, 2015,204(1):163-177.
[13]	DIAZ A, MARTIN-HERNANDEZ A M, DOLCET-SANJUAN R, GARCES-CLAVER A, ALVAREZ J M, GARCIA-MAS J, PICO B, MONFORTE J . Quantitative trait loci analysis of melon (Cucumis melo L.) domestication related traits. Theoretical and Applied Genetics, 2017,130(9):1837-1857.
[14]	马双武, 刘君璞 . 甜瓜种质资源描述规范和数据标准. 北京: 中国农业出版社, 2006.
	MA S W, LIU J P. Descriptors and Data Standard for Melon (Cucumis melo L.). Beijing: China Agriculture Press, 2006. ( in Chinese)
[15]	LUAN F S, DELANNAY I, STAUB J E . Chinese melon ( Cucumis melo L.) diversity analyses provide strategies for germplasm curation, genetic improvement, and evidentiary support of domestication patterns. Euphytica, 2008,164(2):445-461.
[16]	THIEL T, KOTA R, GROSSE I, STEIN N, GRANER A . SNP2CAPS: A SNP and INDEL analysis tool for CAPS marker development. Nucleic Acids Research, 2004,32(1):e5. doi: 10.1093/nar/gnh006
[17]	束永俊, 李勇, 吴娜拉胡, 柏锡, 才华, 纪巍, 朱延明 . 大豆EST-SNP的挖掘、鉴定及其CAPS标记的开发. 作物学报, 2010,36(4):574-579. doi: 10.3724/SP.J.1006.2010.00574
	SHU Y J, LI Y, WU N L H, BAI X, CAI H, JI W, ZHU Y M . Mining and identification of SNP from EST sequences and convertion of CAPS markers in soybean. Acta Agronomica Sinica, 2010,36(4):574-579. (in Chinese) doi: 10.3724/SP.J.1006.2010.00574
[18]	SANDLIN K, PROTHRO J, HEESACKER A, KHALILIAN N, OKASHAH R, XIANG W W, BACHLAVA E, CALDWELL D G, MTAYLOR C A, SEYMOUR D K, WHITE V, CHAN E, TOLLA G, WHITE C, SAFRAN D, GRAHAM E, KNAPP S, MCGREGOR C . Comparative mapping in watermelon [Citrullus lanatus(Thunb.) Matsum.et Nakai]. Tag Theoretical & Applied Genetics Theoretische Und Anbewandte Genetik, 2012,125(8):1603-1618.
[19]	SEBASTIAN P, SCHAEFER H, TELFORD I R H, RENNER S S . Cucumber (Cucumis sativus) and melon(C. melo) have numerous wild relatives in Asia and Australia, and the sister species of melon is from Australia. Proceeding of National Academy of Sciences of the United States of America , 2010,107(32):14269-14273.
[20]	MICHEL P . Phenotypic diversity in wild and cultivated melons (Cucumis melo L). Plant Biotechnology, 2013,30(3):273-278.
[21]	GANAL M W, ALTMANN T, RODER M S . SNP identification in crop plants. Current Opinion in Plant Biology, 2009,12(2):211-217. doi: 10.1016/j.pbi.2008.12.009
[22]	周慧文, 卢丙洋, 马鸿艳, 高鹏, 栾非时, 高启帆, 齐国安 . 西瓜种子大小形状相关QTL分析. 园艺学报, 2016,43(4):715-723.
	ZHOU H W, LU B Y, MA H Y, GAO P, LUAN F S, GAO Q F, QI G A . QTL mapping of watermelon seed traits. Acta Horticulturae Sinica, 2016,43(4):715-723. (in Chinese)
[23]	程瑶, 刘识, 朱子成, 栾非时 . 西瓜糖含量相关性状的QTL分析. 中国蔬菜, 2016(2):24-31.
	CHENG Y, LIU S, ZHU Z C, LUAN F S . QTL analysis for sugar content related traits in watermelon. China Vegetables, 2016(2):24-31. (in Chinese)
[24]	乔军, 陈钰辉, 王利英, 刘富中, 张映, 连勇 . 茄子果形的QTL定位. 园艺学报, 2012,39(6):1115-1122.
	QIAO J, CHEN Y H, WANG L Y, LIU F Z, ZHANG Y, LIAN Y . QTL analysis for fruit shape in eggplant based on genetic linkage map. Acta Horticulturae Sinica, 2012,39(6):1115-1122. (in Chinese)
[25]	TENG W, LI W, ZHANG Q, WU D, ZHAO X, LI H, HAN Y, LI W . Identification of quantitative trait loci underlying seed protein content of soybean including main, epistatic and QTL × environment effects in different regions of Northeast China. Crop and Pasture Science, 2017,68(8):649-655.
[26]	REN Y, MCGREGOR C, ZHANG Y, GONG G Y, ZHANG H Y, GUO S G, SUN H H, CAI W T, ZHANG J, XU Y . An integrated genetic map based on four mapping populations and quantitative trait loci associated with economically important traits in watermelon (Curullus lanatus). BMC Plant Biology, 2014,14(1):33.
[27]	赵璞, 刘瑞响, 李成璞, 刑向茹, 曹晓良, 陶勇生, 张祖新 . 基于掖478导人系的玉米产量性状QTL鉴定. 中国农业科学, 2011,44(17):3508-3519. doi: 10.3864/j.issn.0578-1752.2011.17.003
	ZHAO P, LIU R X, LI C P, XING X R, CAO X L, TAO Y S, ZHANG Z X . QTL mapping for grain yield associated traits using Ye 478 introgression lines in maize. Scientia Agricultura Sinica, 2011,44(17):3508-3519. (in Chinese) doi: 10.3864/j.issn.0578-1752.2011.17.003
[28]	CHAIM A B, PARAN I, GRUBR R C, JAHN M, WIJK R V, PELEMAN J . QTL mapping of fruit-related traits in pepper (Capsicum annuum). Theoretical & Applied Genetics, 2001,102(6/7):1016-1028.
[29]	SALIBA-COLOMBANI V, CAUSSE M, LANGLOIS D, PHILOUZE J, BURET M . Genetic analysis of organoleptic quality in fresh market tomato. 1. Mapping QTLs for physical and chemical traits. Theoretical & Applied Genetics, 2001,102(2/3):259-272.
[30]	陈美霞, 祁建民, 方平平, 李爱青, 危成林, 陶爱芬, 徐建堂, 谢增荣, 林培青, 兰涛, 吴建梅, 陈福成 . 红麻6个重要产量性状的QTL定位. 中国农业科学, 2011,44(5):874-883.
	CHEN M X, QI J M, FANG P P, LI A Q, WEI C L, TAO A F, XU J T, XIE Z R, LIN P Q, LAN T, WU J M, CHEN F C . QTL mapping of six yield traits in kenaf. Scientia Agricultura Sinica, 2011,44(5):874-883. (in Chinese)
[31]	栾非时, 矫士琦, 盛云燕, 朱子成 . 甜瓜果实相关性状QTL分析. 东北农业大学学报, 2017,48(3):1-9.
	LUAN F S, JIAO S Q, SHENG Y Y, ZHU Z C . Mapping of QTL for fruit traits in melon. Journal of Northeast Agricultural University, 2017,48(3):1-9. (in Chinese)
[32]	WANG Y H, WU D H, HUANG J H, TSAO S J, HWU K K, LO H F . Mapping quantitative trait loci for fruit traits and powdery mildew resistance in melon (Cucumis melo). Botanical Studies, 2016,57(1):19.
[33]	栾非时, 卢丙洋, 周慧文, 王学征 . 甜瓜遗传图谱构建及果实相关性状QTL定位. 东北农业大学学报, 2016,47(8):9-20.
	LUAN F S, LU B Y, ZHOU H W, WANG X Z . Construction of genetic linkage map and QTL location for fruit-related traits in melon. Journal of Northeast Agricultural University, 2016,47(8):9-20. (in Chinese)

性状 Trait	亲本及F₁均值 The mean value of and F₁ generation			F_2:3群体 Distribution in F_2:3 population
性状 Trait	母本 M4-130	父本 X207	F₁	均值 Mean	标准差 SD	范围 Range	峰度 SEK	偏度 SES
单果重 Fruit weight (g)	833.0±7.64	21.67±2.87	226.67±50.58	155.92	8.76	33.00—590.00	6.57	2.08
果实长度 Fruit length (cm)	14.20±0.40	3.63±0.23	9.83±1.27	7.87	1.92	3.80—13.30	0.09	0.43
果实宽度 Fruit width (cm)	11.40±0.20	3.30±0.17	6.57±0.31	6.01	1.42	3.00—11.00	0.79	0.74
果肉厚度 Fruit thickness (cm)	3.10±0.10	0.33±0.06	1.13±0.15	1.16	0.34	0.30—2.30	0.58	0.41
果形指数 Fruit shape index (%)	1.25±0.01	1.10±0.12	1.49±0.12	1.32	0.22	0.90—2.10	0.65	0.72
可溶性固形物含量Soluble solids (%)	8.17±0.35	10.5±0.79	10.00±2.20	9.46	1.81	5.90—14.40	-0.55	0.30

性状 Trait	单果重 Fruit weight	果实长度 Fruit length	果实宽度 Fruit width	果肉厚度 Fruit thickness	果形指数 Fruit shape	可溶性固形物含量 Soluble solids
单果重 Fruit weight	1
果实长度 Fruit length	0.873**	1
果实宽度 Fruit width	0.891**	0.832**	1
果肉厚度 Fruit thickness	0.866**	0.813**	0.854**	1
果形指数 Fruit shape	-0.018	0.284**	-0.278**	-0.065	1
可溶性固形物含量 Soluble solids	0.745**	0.706**	0.685**	0.648**	0.034	1

染色体 Chr.	标记数目 No. of Marker	长度 Genetic distance of linkage groups (cM)	标记间平均距离 Average distance of linkage groups (cM)
1	13	91.53	7.04
2	13	112.55	8.66
3	19	121.62	6.40
4	17	140.25	8.25
5	15	108.46	7.23
6	15	160.37	10.69
7	15	166.20	11.08
8	15	144.30	9.62
9	14	155.56	11.11
10	20	107.28	5.36
11	14	163.11	11.65
12	15	129.22	8.61
总计Total	185	1600.45	8.65

通路名称 Pathway	通路注释 Description of pathway	基因数目 Gene number	P值 P-value	基因 Gene
PWY-6902	几丁质降解II Chitin degradation II	6	3.70E-08	MELO3C007961, MELO3C007962, MELO3C007963, MELO3C007964, MELO3C007965, MELO3C007966
PWY-3561	胆碱生物合成III Choline biosynthesis III	2	0.0112	MELO3C007944, MELO3C007945
ASPARAGINE-DEG1-PWY	L-天冬胺酰降解I L-asparagine degradation I	1	0.0502	MELO3C008013
GLYSYN-ALA-PWY	甘氨酸生物合成III Glycine biosynthesis III	1	0.0403	MELO3C014568
PWY-6013	克螨酯生物合成Crepenynate biosynthesis	1	0.0204	MELO3C009630
PWY-6515	根皮苷生物合成 Phloridzin biosynthesis	1	0.0502	MELO3C007921
PWY-7039	磷脂代谢 Phosphatidate metabolism	2	0.0466	MELO3C007944, MELO3C007945