香蕉GDSL脂肪酶基因家族全基因组鉴定与表达分析

张城瑜; 任纹慧; 耿小慧; 李丹; 吴昊; 倪珊珊; 王梦鸽; 罗彬彬; 徐涵; 赖钟雄

doi:10.19303/j.issn.1008-0384.2022.011.007

香蕉GDSL脂肪酶基因家族全基因组鉴定与表达分析

doi: 10.19303/j.issn.1008-0384.2022.011.007

张城瑜^1,,
任纹慧¹,
耿小慧¹,
李丹¹,
吴昊¹,
倪珊珊¹,
王梦鸽¹,
罗彬彬¹,
徐涵^{1, 2},
赖钟雄^1, ,

1.
福建农林大学园艺学院/园艺植物生物工程研究所，福建　福州　350002
2.
图卢兹综合科学研究所，法国　图卢兹　31300

基金项目: 国家重点研发计划项目（2019YFD1000901）；国家现代农业产业技术体系（香蕉）建设专项（GARS-31-15）；福建省高原学科建设项目（102/71201801101）

详细信息

作者简介:
张城瑜（1996−），男，硕士，研究方向：园艺植物生物技术（E-mail：738293225@qq.com）

通讯作者:
赖钟雄（1966−），男，博士，研究员，研究方向：园艺植物生物技术与遗传资源（E-mail：laizx01@163.com）

中图分类号: S 668.1
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出版历程
- 收稿日期: 2022-03-24
- 修回日期: 2022-06-14
- 网络出版日期: 2022-12-28
- 刊出日期: 2022-11-28

Genome-wide Identification and Expressions of Banana GDSL Lipase Gene Family

1.
Institute of Horticultural Biotechnology, Fujian Agriculture and Forestry University, Fuzhou, Fujian　350002, China
2.
Institute de la Recherche Interdisciplinaire de Toulouse, IRIT-ARI, Toulouse　31300, France

摘要

摘要: 目的揭示香蕉GDSL脂肪酶基因家族（MaGDSL）序列特征及其在香蕉生长发育过程中的潜在功能。方法利用生物信息学方法对MaGDSL进行全基因组鉴定，分析其染色体、启动子顺式作用元件和转录因子结合位点（TFBS）的分布情况以及编码蛋白的理化性质、基因结构、保守基序和系统进化关系，并基于转录组数据库分析MaGDSLs在高温（45 ℃）/低温（4 ℃）处理的叶片、枯萎病菌 FocTR4侵染的根系以及自然成熟/乙烯催熟的不同成熟阶段的果实中的表达模式，同时利用实时荧光定量PCR（qRT-PCR）分析部分MaGDSL成员在花粉中的表达情况。结果香蕉A基因组中有76个MaGDSL成员，分布于11个染色体上，可分为9个亚家族，各成员编码区长度为1014～ 2193 bp，其中有5个成员含有不同数量的转录本；多数MaGDSL成员包含5个外显子和4个内含子，编码蛋白具有信号肽且主要定位在内外膜上；启动子区存在多种植物激素和逆境胁迫相关响应元件以及22种TFBSs；MaGDSLs中有3个串联重复基因簇位于4、8号染色体，6个串联重复基因对分别位于1、6、7、8、9和10号染色体上，22对片段重复基因分布在除11号染色体外的所有染色体上。MaGDSLs在香蕉叶片和根部的表达水平差异较大，且个别成员受高、低温胁迫以及枯萎病菌调控，其中5个成员（MaGDSL4-5、8-1、8-12、9-4、9-5）受高、低温胁迫抑制表达，而MaGDSL2-1和MaGDSL6-8受高、低温胁迫诱导表达，MaGDSL4-1和MaGDSL11-1受低温和FocTR4调控，MaGDSL5～8仅响应低温胁迫，其中MaGDSL2-2和MaGLP10-5对各种处理均有响应，而MaGDSL1-1在根系和花粉中均有高表达。结论 MaGDSLs可能在香蕉生长发育过程中发挥重要作用，同时部分成员特异响应生物和非生物胁迫。
- 香蕉 /
- GDSL脂肪酶 /
- 基因组 /
- 逆境胁迫 /
- 表达分析
Abstract: Objective Sequences and functions of banana GDSL lipase gene family (MaGDSL) were studied. Methods Bioinformatics was applied to identify the genomes, analyze the distributions of chromosomes, promoter cis-acting elements, and transcription factor binding sites (TFBS), and determine the physicochemical properties, gene structure, conserved motifs, and phylogenetic relationships of the encoded proteins of the MaGDSL family. Based on the transcriptome database, the expressions of MaGDSLs in the high (45 ℃)- or low temperature (4 ℃)-treated leaves, FocTR4-infested roots, and the natural or ethylene ripened fruits were measured. And qRT-PCR was employed to obtain the expressions of MaGDSL members in pollen. Results The banana A genome had 76 MaGDSLs of 9 subfamilies distributed on 11 chromosomes. The coding region of each member was 1014–2193 bp. Five of the members contained varied number of transcripts, but most of them had 5 exons and 4 introns. The encoded proteins had signal peptides localized mainly in the inner and outer membranes. The MaGDSLs had 3 tandem repeat clusters on chromosomes 4 and 8, 6 tandem repeat pairs on chromosomes 1, 6, 7, 8, 9, and 10, and 22 fragment repeat pairs on all except chromosome 11. The expressions of MaGDSLs in banana leaves and roots were highly variable with individual members regulated by high and low temperature stresses as well as the blast fungus. Five members (i.e., MaGDSL4-5, 8-1, 8-12, 9-4, 9-5) were repressed, but MaGDSL2-1 and MaGDSL6-8 induced, by high- or low-temperature exposure, while MaGDSL4-1 and MaGDSL11-1 regulated by low temperature and FocTR4, MaGDSL5–8 responded only to low temperature, MaGDSL2-2 and MaGLP10-5 sensitive to various treatments, and MaGDSL1-1 highly expressed in the roots and pollens. Conclusion MaGDSLs might play an important role in the growth and development of banana plants. Some of the members responded specifically to certain biotic and/or abiotic stresses.
- Banana /
- GDSL lipase gene /
- genome /
- stress by adversity /
- expression analysis

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图 1 MaGDSL家族成员的系统进化树

Figure 1. Phylogenetic tree of MaGDSLs in banana

下载: 全尺寸图片幻灯片

图 2 香蕉MaGDSL基因家族的蛋白保守基序及其基因结构分布图

Figure 2. Conserved motifs in encoded proteins and gene structure of MaGDSLs

下载: 全尺寸图片幻灯片

图 3 MaGDSL 的染色体定位和共线性分布

绿色线表示 MaGDSL 的片段复制基因对，标红的基因表示 MaGDSL 的串联复制基因。

Figure 3. Chromosome localization and collinear distribution of MaGDSLs

Green line indicates segmental duplication gene pairs of MaGDSL; red color indicates tandem duplication genes of MaGDSL.

下载: 全尺寸图片幻灯片

图 4 MaGDSL 基因家族启动子顺式作用元件分析

Figure 4. Analysis of cis-acting elements of MaGDSL gene family promoter

下载: 全尺寸图片幻灯片

图 5 MaGDSL 基因家族启动子的转录因子结合位点分布

Figure 5. Distribution of transcription factor binding sites in MaGDSLs promoter

下载: 全尺寸图片幻灯片

图 6 MaGDSL 家族成员表达量热图

Figure 6. Heatmap for expression of MaGDSL gene family members

下载: 全尺寸图片幻灯片

图 7 MaGDSL家族部分成员在福州野生蕉最外层花粉的相对表达量

Figure 7. Relative expressions of MaGDSLs in outermost pollen layer of wild Fuzhou banana plants

下载: 全尺寸图片幻灯片

表 1 qPCR 分析引物

Table 1. Primers for qRT-PCR

基因名称 Gene ID	引物名称 Primer name	引物序列 Primer sequence
内参UBQ	F	3′-GGCACCACAAACAACACAGG-5′
内参UBQ	R	3′-AGACGAGCAAGGCTTCCATT-5′
MaGDSL2-7	F	3′-GGAAGACGGGAGAGGGATAC-5′
MaGDSL2-7	R	3′-GCCTACGTTCATGGGGTAAGC-5′
MaGDSL3-2	F	3′-ATCTTCGGAGACTCCCTCGT-5′
MaGDSL3-2	R	3′-GTGCTTGCCCTCTTGTGTTT-5′
MaGDSL5-1	F	3′-TTGATGCAGCAACAGTGACA-5′
MaGDSL5-1	R	3′-AAACATAGCAGGCACCGACT-5′
MaGDSL6-4	F	3′-GAGCTGAGAAGGACGATGCT-5′
MaGDSL6-4	R	3′-GTCAAGCCATTGGTGAACCT-5′
MaGDSL6-7	F	3′-CCCAATGGCAACAAGTATCC-5′
MaGDSL6-7	R	3′-CCTGAGGGGCTACATACCAA-5′
MaGDSL6-9	F	3′-TTGCCAGGCTCTTTCTTTGT-5′
MaGDSL6-9	R	3′-AGCACTTGTCAGCGATGATG-5′
MaGDSL7-5	F	3′-TGAGGGCTCCATGGACATA-5′
MaGDSL7-5	R	3′-GCAAGTTCACGGGATAGCA-5′
MaGDSL8-10	F	3′-CGGAACCACAGGACGATACT-5′
MaGDSL8-10	R	3′-TGTCCCAAAGGATTCCAGAG-5′
MaGDSL9-2	F	3′-ATGAACTCAACGCCTTGGAC-5′
MaGDSL9-2	R	3′-GATGAGGACGATTGCAGCTT-5′
MaGDSL11-6	F	3′-TGCTTCCCATGCTATCGAG-5′
MaGDSL11-6	R	3′-CCCAAATGGTCTGCAACAG-5′
MaGDSL1-1	F	3′-CGTTGTGATTCTGTGCTGCT-5′
MaGDSL1-1	R	3′-GTTTCTCAGCGAGGAAGTCG-5′

下载: 导出CSV

表 2 MaGDSL家族成员基本信息和编码蛋白理化性质

Table 2. Information and physiochemical properties of MaGDSL encoded proteins

基因 ID Gene ID	染色体位置 Chromosome location	基因 Gene	长度 Length/aa	等电点 pI	不稳定指数 Instability index	亲水性 GRAVY	信号肽 Signal peptide	跨膜结构 Transmembrane domain	亚细胞定位 Protein subcellular localization Prediction
Ma01_g03770	chr01:2506772..2513988(−)	MaGDSL1-1	362	5.97	40.6	−0.118	是	0	外膜 Outer membrane
Ma01_g03780	chr01:2514735..2517040(+)	MaGDSL1-2	363	5.58	36.14	−0.064	是	0	胞外 Extracellular
Ma01_g04800	chr01:3286719..3288349(−)	MaGDSL1-3	363	6.90	39.64	0.034	是	1	外膜 Outer membrane
Ma01_g17620	chr01:12898520..12900562(+)	MaGDSL1-4	390	6.25	42.89	−0.045	是	0	外膜 Outer membrane
Ma02_g12980	chr02:21545341..21546931(+)	MaGDSL2-1	355	6.52	32.18	0.092	是	1	周质 Periplasmic
Ma02_g13020	chr02:21592310..21594060(+)	MaGDSL2-2	355	6.22	33.06	0.084	是	1	外膜 Outer membrane
Ma02_g13050	chr02:21608508..21610184(+)	MaGDSL2-3	353	6.99	28.54	0.041	是	1	外膜 Outer membrane
Ma02_g13620	chr02:21929576..21940378(+)	MaGDSL2-4	430	8.05	36.67	−0.026	是	1	外膜 Outer membrane
Ma02_g14360	chr02:22465764..22467578(−)	MaGDSL2-5a	401	5.25	33.59	0.210	否	0	外膜 Outer membrane
Ma02_g14360	chr02:22465764..22467578(−)	MaGDSL2-5b	396	5.24	35.01	0.211	否	0	外膜 Outer membrane
Ma02_g18570	chr02:25021259..25022978(−)	MaGDSL2-6	358	6.23	22.58	0.000	是	0	外膜 Outer membrane
Ma02_g21270	chr02:26951434..26952967(+)	MaGDSL2-7	376	9.21	36.69	−0.090	是	0	外膜 Outer membrane
Ma03_g03330	chr03:2232832..2235731(−)	MaGDSL3-1	419	7.16	32.55	−0.113	否	0	周质 Periplasmic
Ma03_g16970	chr03:22263933..22266432(+)	MaGDSL3-2	366	5.97	35.88	−0.135	是	0	外膜 Outer membrane
Ma03_g25700	chr03:29665153..29668011(+)	MaGDSL3-3a	372	5.79	45.25	0.097	是	0	外膜 Outer membrane
Ma03_g25700	chr03:29665153..29668011(+)	MaGDSL3-3b	364	5.90	47.02	0.081	是	0	周质 Periplasmic
Ma03_g26380	chr03:30141639..30143572(−)	MaGDSL3-4	410	6.35	40.42	−0.079	是	0	外膜 Outer membrane
Ma04_g05070	chr04:3833650..3836264(−)	MaGDSL4-1	356	8.59	34.73	0.044	是	0	胞外 Extracellular
Ma04_g18170	chr04:19831827..19833593(−)	MaGDSL4-2	347	6.64	27.42	0.090	是	0	外膜 Outer membrane
Ma04_g19770	chr04:22407980..22409457(+)	MaGDSL4-3	373	6.09	37.18	0.008	是	0	外膜 Outer membrane
Ma04_g32080	chr04:32208763..32212100(+)	MaGDSL4-4	363	8.25	22.57	0.243	是	0	外膜 Outer membrane
Ma04_g32090	chr04:32217091..32219380(+)	MaGDSL4-5	367	5.13	32.77	0.094	是	0	外膜 Outer membrane
Ma04_g32250	chr04:32308311..32310571(+)	MaGDSL4-6	362	8.54	22.38	0.167	是	0	外膜 Outer membrane
Ma04_g32390	chr04:32366386..32368477(−)	MaGDSL4-7	364	9.47	37.07	−0.018	是	0	外膜 Outer membrane
Ma04_g35750	chr04:34368316..34369739(−)	MaGDSL4-8	365	5.27	43.48	0.075	是	0	细胞质 Cytoplasmic
Ma04_g37620	chr04:35437527..35439771(+)	MaGDSL4-9a	358	8.75	37.23	0.019	是	0	外膜 Outer membrane
Ma04_g37620	chr04:35437527..35439771(+)	MaGDSL4-9b	358	8.75	37.23	0.019	是	0	外膜 Outer membrane
Ma05_g01340	chr05:780195..783192(−)	MaGDSL5-1	413	5.28	20.21	−0.015	是	0	胞外 Extracellular
Ma05_g10980	chr05:7975525..7978280(+)	MaGDSL5-2	355	6.03	40.10	−0.082	是	0	外膜 Outer membrane
Ma05_g14230	chr05:10339954..10341215(+)	MaGDSL5-3	361	5.35	37.32	−0.055	是	0	细胞 Cytoplasmic
Ma05_g15830	chr05:12426846..12430706(+)	MaGDSL5-4	375	9.05	39.01	−0.130	是	1	胞外 Extracellular
Ma05_g17280	chr05:19801910..19803428(−)	MaGDSL5-5	366	6.12	38.93	−0.015	是	1	外膜 Outer membrane
Ma05_g17650	chr05:21042672..21044837(+)	MaGDSL5-6	386	8.71	36.61	−0.085	是	0	周质 Periplasmic
Ma05_g23460	chr05:35527091..35528767(−)	MaGDSL5-7	374	6.75	35.27	−0.146	是	1	外膜 Outer membrane
Ma05_g23660	chr05:35833232..35837552(+)	MaGDSL5-8	377	5.49	43.04	−0.016	是	0	外膜 Outer membrane
Ma06_g04310	chr06:3104879..3106481(−)	MaGDSL6-1a	372	5.40	31.37	−0.102	是	0	外膜 Outer membrane
Ma06_g04310	chr06:3104879..3106481(−)	MaGDSL6-1b	338	5.52	30.79	0.033	是	0	外膜 Outer membrane
Ma06_g24960	chr06:25117515..25118735(−)	MaGDSL6-2	355	6.79	34.76	0.092	是	1	内膜 Inner membrane
Ma06_g25030	chr06:25237635..25243333(−)	MaGDSL6-3	380	8.23	37.33	0.008	是	0	周质 Periplasmic
Ma06_g27800	chr06:29688900..29691154(−)	MaGDSL6-4	365	5.20	42.08	−0.121	是	0	外膜 Outer membrane
Ma06_g29860	chr06:31225349..31227058(−)	MaGDSL6-5	411	7.56	34.71	−0.013	否	0	周质 Periplasmic
Ma06_g30580	chr06:31830895..31832818(+)	MaGDSL6-6a	375	8.97	43.59	0.022	是	0	周质 Periplasmic
Ma06_g30580	chr06:31830895..31832818(+)	MaGDSL6-6b	374	8.86	43.71	0.032	是	0	周质 Periplasmic
Ma06_g30640	chr06:31860762..31862418(−)	MaGDSL6-7	447	9.21	31.85	−0.149	否	0	外膜 Outer membrane
Ma06_g37580	chr06:36585560..36587436(−)	MaGDSL6-8	398	5.94	41.69	−0.055	否	1	胞外 Extracellular
Ma06_g38100	chr06:36975407..36978177(+)	MaGDSL6-9	365	6.50	42.06	0.061	是	0	内膜 Inner membrane
Ma07_g08760	chr07:6557127..6558543(+)	MaGDSL7-1	359	7.59	24.74	−0.001	是	0	外膜 Outer membrane
Ma07_g12320	chr07:9208788..9210632(+)	MaGDSL7-2	389	6.06	30.74	0.000	否	1	外膜 Outer membrane
Ma07_g13910	chr07:10418714..10420475(+)	MaGDSL7-3	360	8.08	31.58	0.016	是	0	内膜 Inner membrane
Ma07_g13920	chr07:10420620..10422694(+)	MaGDSL7-4	365	9.53	42.01	−0.077	是	0	外膜 Outer membrane
Ma07_g14440	chr07:10888114..10890751(+)	MaGDSL7-5	363	5.12	49.96	−0.009	否	0	外膜 Outer membrane
Ma08_g06100	chr08:4121189..4123492(−)	MaGDSL8-1	371	8.55	37.99	−0.007	是	0	周质 Periplasmic
Ma08_g09750	chr08:7078632..7081137(+)	MaGDSL8-2	366	6.60	31.02	0.054	是	0	外膜 Outer membrane
Ma08_g10150	chr08:7392254..7396118(−)	MaGDSL8-3	381	6.65	33.87	−0.001	是	0	外膜 Outer membrane
Ma08_g10160	chr08:7397273..7398935(−)	MaGDSL8-4	404	6.54	37.62	−0.101	否	0	周质 Periplasmic
Ma08_g10170	chr08:7404987..7406648(−)	MaGDSL8-5	367	6.26	39.19	−0.081	是	0	外膜 Outer membrane
Ma08_g10480	chr08:7659127..7660505(+)	MaGDSL8-6	376	5.36	23.81	0.106	是	1	外膜 Outer membrane
Ma08_g21060	chr08:35064951..35067710(+)	MaGDSL8-7	359	6.88	31.43	−0.068	是	0	内膜 Inner membrane
Ma08_g21070	chr08:35068690..35077978(+)	MaGDSL8-8	731	8.61	38.45	0.113	是	0	外膜 Outer membrane
Ma08_g28640	chr08:40487336..40489392(−)	MaGDSL8-9	384	5.02	45.15	−0.050	是	1	胞外 Extracellular
Ma08_g30430	chr08:41700756..41702316(+)	MaGDSL8-10	397	6.70	39.81	0.115	是	0	内膜 Inner membrane
Ma08_g30470	chr08:41727515..41729409(+)	MaGDSL8-11	356	5.16	31.84	0.142	是	0	细胞外 Extracellular
Ma08_g30480	chr08:41730080..41733085(+)	MaGDSL8-12	383	4.54	39.03	0.116	是	1	外膜 Outer membrane
Ma09_g01870	chr09:1401507..1405717(+)	MaGDSL9-1	381	7.11	35.50	−0.111	是	0	周质 Periplasmic
Ma09_g05540	chr09:3558484..3560119(−)	MaGDSL9-2	369	6.89	33.29	−0.049	是	0	外膜 Outer membrane
Ma09_g19240	chr09:21599052..21600910(−)	MaGDSL9-3	365	4.94	42.59	0.050	是	1	内膜 Inner membrane
Ma09_g24520	chr09:36179911..36182176(+)	MaGDSL9-4	360	7.61	35.81	0.074	是	1	胞外 Extracellular
Ma09_g24530	chr09:36193365..36196041(+)	MaGDSL9-5	360	8.73	35.21	0.077	是	0	胞外 Extracellular
Ma10_g12640	chr10:25677743..25681254(+)	MaGDSL10-1	381	6.16	38.28	0.108	是	1	周质 Periplasmic
Ma10_g14470	chr10:26912910..26916293(+)	MaGDSL10-2	360	6.59	28.16	−1.019	是	0	外膜 Outer membrane
Ma10_g14480	chr10:26917998..26919683(−)	MaGDSL10-3	370	7.98	42.03	0.025	是	0	周质 Periplasmic
Ma10_g14730	chr10:27073351..27075394(+)	MaGDSL10-4	393	7.63	39.37	0.040	是	0	周质 Periplasmic
Ma10_g21620	chr10:31338004..31344227(+)	MaGDSL10-5	359	7.23	33.11	0.023	是	0	内膜 Inner membrane
Ma11_g05100	chr11:3940477..3942390(+)	MaGDSL11-1	374	5.36	34.01	0.059	是	0	外膜 Outer membrane
Ma11_g09200	chr11:7355875..7357479(−)	MaGDSL11-2	374	5.28	34.44	0.075	是	0	周质 Periplasmic
Ma11_g10620	chr11:10119577..10121852(−)	MaGDSL11-3	366	8.87	34.88	−0.042	是	0	胞外 Extracellular
Ma11_g14690	chr11:20398154..20399698(+)	MaGDSL11-4	368	7.50	37.23	−0.133	是	0	外膜 Outer membrane
Ma11_g15240	chr11:20938047..20940295(−)	MaGDSL11-5	360	6.01	37.08	0.070	是	0	外膜 Outer membrane
Ma11_g16030	chr11:21634731..21637078(−)	MaGDSL11-6	361	5.49	32.42	−0.033	是	0	胞外 Extracellular
Ma11_g18800	chr11:23822530..23824109(+)	MaGDSL11-7	367	5.34	28.72	−0.015	是	1	外膜 Outer membrane
Ma11_g25050	chr11:27802837..27804653(+)	MaGDSL11-8	386	6.49	31.58	−0.006	是	1	周质 Periplasmic
跨膜结构中，0代表不存在跨膜结构，1代表存在跨膜结构。 In the table, 0 means no transmembrane domain existed, 1 transmembrane domain existed.

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表 3 MaGDSLs同源基因 Ka/Ks 分析

Table 3. Ka/Ks analysis of MaGDSLs homologous genes

复制类型 Duplication type	基因名称 Gene name	基因ID Gene ID	基因名称 Gene name	基因ID Gene ID	Ka	Ks	Ka/Ks
串联重复 Tandem duplication	MaGDSL1-1	Ma01_g03770	MaGDSL1-2	Ma01_g03780	0.0567	0.1592	0.3562
	MaGDSL4-4	Ma04_g32080	MaGDSL4-5	Ma04_g32090	0.3667	1.5348	0.2389
	MaGDSL4-6	Ma04_g32250	MaGDSL4-7	Ma04_g32390	0.7590	2.1143	0.3590
	MaGDSL6-6	Ma06_g30580	MaGDSL6-7	Ma06_g30640	0.9792	1.6565	0.5911
	MaGDSL7-3	Ma07_g13910	MaGDSL7-4	Ma07_g13920	0.3650	1.2292	0.2969
	MaGDSL8-3	Ma08_g10150	MaGDSL8-4	Ma08_g10160	0.4687	1.4135	0.3316
	MaGDSL8-4	Ma08_g10160	MaGDSL8-5	Ma08_g10170	0.0355	0.0230	1.5416
	MaGDSL8-7	Ma08_g21060	MaGDSL8-8	Ma08_g21070	0.2879	1.1093	0.2596
	MaGDSL8-10	Ma08_g30430	MaGDSL8-11	Ma08_g30470	0.8306	——	——
	MaGDSL8-11	Ma08_g30470	MaGDSL8-12	Ma08_g30480	0.4536	1.4482	0.3132
	MaGDSL9-4	Ma09_g24520	MaGDSL9-5	Ma09_g24530	0.0225	0.0303	0.7449
	MaGDSL10-2	Ma10_g14470	MaGDSL10-3	Ma10_g14480	0.4506	1.3564	0.3322
片段复制 Segmental duplication	MaGDSL1-3	Ma01_g04800	MaGDSL5-5	Ma05_g17280	0.0963	0.7646	0.1259
	MaGDSL2-4	Ma02_g13620	MaGDSL4-8	Ma04_g35750	0.1904	0.7111	0.2678
	MaGDSL2-6	Ma02_g18570	MaGDSL7-1	Ma07_g08760	0.1022	0.5938	0.1721
	MaGDSL3-3	Ma03_g25700	MaGDSL8-1	Ma08_g06100	0.1140	0.5042	0.2262
	MaGDSL3-2	Ma03_g16970	MaGDSL9-2	Ma09_g05540	0.1436	0.7192	0.1997
	MaGDSL3-1	Ma03_g03330	MaGDSL10-2	Ma10_g14470	0.1048	0.5944	0.1764
	MaGDSL4-4	Ma04_g32080	MaGDSL5-2	Ma05_g10980	0.3694	2.3606	0.1565
	MaGDSL4-1	Ma04_g05070	MaGDSL9-4	Ma09_g24520	0.1954	0.6881	0.2840
	MaGDSL5-8	Ma05_g23660	MaGDSL8-3	Ma08_g10150	0.1251	0.4943	0.2531
	MaGDSL5-6	Ma05_g17650	MaGDSL9-1	Ma09_g01870	0.1104	0.6648	0.1661
	MaGDSL5-8	Ma05_g23660	MaGDSL11-1	Ma11_g05100	0.4539	1.8844	0.2409
	MaGDSL6-4	Ma06_g27800	MaGDSL9-2	Ma09_g05540	0.0962	0.5062	0.1901
	MaGDSL7-3	Ma07_g13910	MaGDSL8-7	Ma08_g21060	0.1133	0.4296	0.2638
	MaGDSL7-4	Ma07_g13920	MaGDSL8-8	Ma08_g21070	0.2226	1.3169	0.1691
	MaGDSL7-3	Ma07_g13910	MaGDSL10-5	Ma10_g21620	0.0939	0.4276	0.2196
	MaGDSL7-3	Ma07_g13910	MaGDSL10-2	Ma10_g14470	0.2573	1.1430	0.2251
	MaGDSL7-4	Ma07_g13920	MaGDSL10-3	Ma10_g14480	0.4068	1.3864	0.2934
	MaGDSL8-7	Ma08_g21060	MaGDSL10-5	Ma10_g21620	0.0923	0.5003	0.1846
	MaGDSL8-2	Ma08_g09750	MaGDSL11-7	Ma11_g18800	0.1081	0.7247	0.1491
	MaGDSL8-4	Ma08_g10160	MaGDSL11-1	Ma11_g05100	0.3611	1.0690	0.3378
	MaGDSL8-11	Ma08_g30470	MaGDSL11-3	Ma11_g10620	0.2212	0.5409	0.4090
	MaGDSL10-2	Ma10_g14470	MaGDSL10-5	Ma10_g21620	0.2361	0.9899	0.2385

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