Genetic diversity analysis and fingerprint construction of Dading bitter gourd germplasm resources

doi:10.3969/j.issn.1004-1524.20230063

Acta Agriculturae Zhejiangensis ›› 2024, Vol. 36 ›› Issue (1): 103-114.DOI: 10.3969/j.issn.1004-1524.20230063

• Horticultural Science • Previous Articles Next Articles

Genetic diversity analysis and fingerprint construction of Dading bitter gourd germplasm resources

YANG Tianwen¹(), WANG Jing¹, LI Jiong¹, XU Binqi¹, CHENG Jiaowen², HONG Yu¹, CAO Yi¹, CUI Junjie¹^,^*()

1. Department of Horticulture, Foshan University, Foshan 528225, Guangdong, China
2. College of Horticulture, South China Agricultural University, Guangzhou 510642, China

Received:2023-01-18 Online:2024-01-25 Published:2024-02-18
Contact: CUI Junjie

Abstract

Abstract:

To achieve rapid and accurate identification of the germplasm resources of the Dading bitter gourd, 26 Dading bitter gourd germplasm resources including the main cultivars were selected as materials, cluster analysis and fingerprint construction were carried out by screening the high polymorphism simple sequence repeat (SSR) core markers. The results showed that among the 6 SSR motif unit types in the bitter gourd genome, the polymorphism of dinucleotide SSR markers was the best, which was suitable for further screening of SSR core markers. At the genetic similarity coefficient of 0.80, 40 SSR markers clustered the 26 Dading bitter gourd germplasm resources into 3 main groups. The first group contained 18 germplasm resources, and the fruit shape was mainly short conical. The second group contained 5 germplasm resources, and the fruit shape was mainly long conical. The third group contained 3 germplasm resources, consisting of materials with fruits similar to those of Dading bitter gourds but containing non-Dading bitter gourd germplasm backgrounds. The 4 SSR core markers, MC05_69594, MC04_50530, MC06_87314 and MC10_146038, were used to effectively identify 26 Dading bitter gourd germplasm resources, and a unique QR code fingerprint was established for each Dading bitter gourd germplasm, among which No. 1 and No. 3 were suspected to be similar materials. Taken together, the fingerprint of Dading bitter gourd could provide a scientific basis for the germplasm identification and breeding utilization for Dading bitter gourd.

Key words: dading bitter gourd, simple sequence repeat marker, genetic diversity, germplasm identification, fingerprinting

CLC Number:

S642.5

YANG Tianwen, WANG Jing, LI Jiong, XU Binqi, CHENG Jiaowen, HONG Yu, CAO Yi, CUI Junjie. Genetic diversity analysis and fingerprint construction of Dading bitter gourd germplasm resources[J]. Acta Agriculturae Zhejiangensis, 2024, 36(1): 103-114.

Figures/Tables 9

References 28

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编号 Number	材料名称 Material name	种类 Species	种质类型 Germplasm type	来源 Origin	果实部分特征 Some characteristics of fruit
1	东来大顶 Donglai	M. charantia	杂交种 Hybrid	百蔬网	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
2	1号大顶 NO.1	M. charantia	杂交种 Hybrid	百蔬网	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
3	3号大顶 NO.3	M. charantia	杂交种 Hybrid	百蔬网	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
4	翠绿三号 Cuilyu NO.3	M. charantia	杂交种 Hybrid	广东科农蔬菜种业有限公司 Guangdong Kenong Vegetable Seed Company	长圆锥形,绿色果皮,黄籽 Long conical, green peel, yellow seed coat
5	清蔬2号 Qingshu NO.2	M. charantia	杂交种 Hybrid	清远市清蔬种子有限公司 Qingyuan Qingshu Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
6	金利塔 Jinlita	M. charantia	杂交种 Hybrid	清远市清蔬种子有限公司 Qingyuan Qingshu Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
7	巨顶大顶 Juding	M. charantia	杂交种 Hybrid	广州市义农园艺有限公司 Guangzhou Yinong Horticulture Company	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
8	珍珠大顶 Zhenzhu	M. charantia	杂交种 Hybrid	台山市利农种子有限公司 Taishan Linong Seed Company	长圆锥形,绿色果皮,黄籽 Long conical, green peel, yellow seed coat
9	超绿大顶 Chaolyu	M. charantia	杂交种 Hybrid	香港惟勤 Hong Kong Weqin	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
10	科丰大顶 Kefeng	M. charantia	杂交种 Hybrid	广州科田种苗有限公司 Guangzhou Ketian Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
11	利农珍珠 LNzhenzhu	M. charantia	杂交种 Hybrid	广东和利农种业股份有限公司 Guangdong Helinong Seed Company	长圆锥形,浅绿色果皮,黑籽 Long conical, light green peel, black seed coat
12	博优特804 BYT804	M. charantia	杂交种 Hybrid	广州博优特农业科技有限公司 Guangzhou Boyoute Agricultural Technology Company	长圆锥形,绿色果皮,黑籽 Long conical, green peel, black seed coat
13	翠绿厚肉 Cuilyu HR	M. charantia	杂交种 Hybrid	珠海市斗门区井岸兴隆蔬菜种子点 Zhuhai Doumen Jinganxinglong Vegetable Seed Store	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
14	金顶3号 Jinding NO.3	M. charantia	杂交种 Hybrid	金韩种业 Jinhan Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
15	华鼎3号 Huading NO.3	M. charantia	杂交种 Hybrid	华南农业大学华农大种业 South China Agricultural University Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
16	真功夫13号 Zhengongfu NO.13	M. charantia	杂交种 Hybrid	广东和利农种业股份有限公司 Guangdong Helinong Seed Company	长圆锥形,浅绿色果皮,黑籽 Long conical, light green peel, black seed coat
17	特大大顶 Teda	M. charantia	杂交种 Hybrid	金叶子园艺馆 Jinyezi Horticulture House	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
18	青蕾1号 Qinglei NO.1	M. charantia	杂交种 Hybrid	广州乾农农业科技发展有限公司 Guangzhou Gannong Agricultural Technology Company	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
19	美鼎8号 Meiding NO.8	M. charantia	杂交种 Hybrid	揭阳市农研种业有限公司 Jieyang Nongyan Seed Company	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
20	江门大顶 Jiangmen	M. charantia	常规种 Conventional cultivar	子丰种业 Zifeng Seed Company	长圆锥形,绿色果皮,黄籽 Long conical, green peel, yellow seed coat
21	鼎胜大顶 Dingsheng	M. charantia	杂交种 Hybrid	禾之硕农业 Hezhishuo Agriculture	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
22	谭边大顶 Tanbian	M. charantia	地方种 Landrace	收集于广东省佛山市南海区谭边村 Selected from Foshan Tanbian village	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
23	广良2号 Guangliang NO.2	M. charantia	杂交种 Hybrid	子丰种业 Zifeng Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
24	江门F Jiangmen F	M. charantia	自交系 Inbred line	选自江门大顶苦瓜 Selected from Jiangmen Dading bitter gourd	长圆锥形,绿色果皮,黄籽 Long conical, green peel, black seed coat
25	惠州F Huizhou F	M. charantia	自交系 Inbred line	选自惠州大顶苦瓜 Selected from Huizhou Dading bitter gourd	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
26	苹果大顶 Pingguo	M. charantia	常规种 Conventional cultivar	山东寿光欣欣然园艺有限公司 Shouguang Xinxinran Horticulture Company	短或长圆锥形,白色果皮,黄籽 Short or long conical, white peel, yellow seed coat
27	汾溪L Fenxi L	M. charantia	地方种 Landrace	收集于安徽省黄山市祁门县 Selected from Qimen County, Huangshan	椭球形,绿色果皮,黄籽 Spheroid, green peel, yellow seed coat
28	南靖TR Nanjing TR	M. charantia ssp. macroloba	野生亚种 Wild subspecies	收集于福建省漳州市南靖县 Selected from Nanjing County, Zhangzhou	纺锤形,绿色果皮,黑籽 Spindle-shaped, green peel, black seed coat

编号 Number	材料名称 Material name	种类 Species	种质类型 Germplasm type	来源 Origin	果实部分特征 Some characteristics of fruit
1	东来大顶 Donglai	M. charantia	杂交种 Hybrid	百蔬网	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
2	1号大顶 NO.1	M. charantia	杂交种 Hybrid	百蔬网	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
3	3号大顶 NO.3	M. charantia	杂交种 Hybrid	百蔬网	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
4	翠绿三号 Cuilyu NO.3	M. charantia	杂交种 Hybrid	广东科农蔬菜种业有限公司 Guangdong Kenong Vegetable Seed Company	长圆锥形,绿色果皮,黄籽 Long conical, green peel, yellow seed coat
5	清蔬2号 Qingshu NO.2	M. charantia	杂交种 Hybrid	清远市清蔬种子有限公司 Qingyuan Qingshu Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
6	金利塔 Jinlita	M. charantia	杂交种 Hybrid	清远市清蔬种子有限公司 Qingyuan Qingshu Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
7	巨顶大顶 Juding	M. charantia	杂交种 Hybrid	广州市义农园艺有限公司 Guangzhou Yinong Horticulture Company	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
8	珍珠大顶 Zhenzhu	M. charantia	杂交种 Hybrid	台山市利农种子有限公司 Taishan Linong Seed Company	长圆锥形,绿色果皮,黄籽 Long conical, green peel, yellow seed coat
9	超绿大顶 Chaolyu	M. charantia	杂交种 Hybrid	香港惟勤 Hong Kong Weqin	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
10	科丰大顶 Kefeng	M. charantia	杂交种 Hybrid	广州科田种苗有限公司 Guangzhou Ketian Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
11	利农珍珠 LNzhenzhu	M. charantia	杂交种 Hybrid	广东和利农种业股份有限公司 Guangdong Helinong Seed Company	长圆锥形,浅绿色果皮,黑籽 Long conical, light green peel, black seed coat
12	博优特804 BYT804	M. charantia	杂交种 Hybrid	广州博优特农业科技有限公司 Guangzhou Boyoute Agricultural Technology Company	长圆锥形,绿色果皮,黑籽 Long conical, green peel, black seed coat
13	翠绿厚肉 Cuilyu HR	M. charantia	杂交种 Hybrid	珠海市斗门区井岸兴隆蔬菜种子点 Zhuhai Doumen Jinganxinglong Vegetable Seed Store	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
14	金顶3号 Jinding NO.3	M. charantia	杂交种 Hybrid	金韩种业 Jinhan Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
15	华鼎3号 Huading NO.3	M. charantia	杂交种 Hybrid	华南农业大学华农大种业 South China Agricultural University Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
16	真功夫13号 Zhengongfu NO.13	M. charantia	杂交种 Hybrid	广东和利农种业股份有限公司 Guangdong Helinong Seed Company	长圆锥形,浅绿色果皮,黑籽 Long conical, light green peel, black seed coat
17	特大大顶 Teda	M. charantia	杂交种 Hybrid	金叶子园艺馆 Jinyezi Horticulture House	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
18	青蕾1号 Qinglei NO.1	M. charantia	杂交种 Hybrid	广州乾农农业科技发展有限公司 Guangzhou Gannong Agricultural Technology Company	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
19	美鼎8号 Meiding NO.8	M. charantia	杂交种 Hybrid	揭阳市农研种业有限公司 Jieyang Nongyan Seed Company	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
20	江门大顶 Jiangmen	M. charantia	常规种 Conventional cultivar	子丰种业 Zifeng Seed Company	长圆锥形,绿色果皮,黄籽 Long conical, green peel, yellow seed coat
21	鼎胜大顶 Dingsheng	M. charantia	杂交种 Hybrid	禾之硕农业 Hezhishuo Agriculture	短圆锥形,绿色果皮,黄籽 Short conical, green peel, yellow seed coat
22	谭边大顶 Tanbian	M. charantia	地方种 Landrace	收集于广东省佛山市南海区谭边村 Selected from Foshan Tanbian village	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
23	广良2号 Guangliang NO.2	M. charantia	杂交种 Hybrid	子丰种业 Zifeng Seed Company	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
24	江门F Jiangmen F	M. charantia	自交系 Inbred line	选自江门大顶苦瓜 Selected from Jiangmen Dading bitter gourd	长圆锥形,绿色果皮,黄籽 Long conical, green peel, black seed coat
25	惠州F Huizhou F	M. charantia	自交系 Inbred line	选自惠州大顶苦瓜 Selected from Huizhou Dading bitter gourd	短圆锥形,绿色果皮,黑籽 Short conical, green peel, black seed coat
26	苹果大顶 Pingguo	M. charantia	常规种 Conventional cultivar	山东寿光欣欣然园艺有限公司 Shouguang Xinxinran Horticulture Company	短或长圆锥形,白色果皮,黄籽 Short or long conical, white peel, yellow seed coat
27	汾溪L Fenxi L	M. charantia	地方种 Landrace	收集于安徽省黄山市祁门县 Selected from Qimen County, Huangshan	椭球形,绿色果皮,黄籽 Spheroid, green peel, yellow seed coat
28	南靖TR Nanjing TR	M. charantia ssp. macroloba	野生亚种 Wild subspecies	收集于福建省漳州市南靖县 Selected from Nanjing County, Zhangzhou	纺锤形,绿色果皮,黑籽 Spindle-shaped, green peel, black seed coat

SSR类型 SSR type	数量 Number	扩增引物数量 Number of amplified marker	扩增率 Amplification rate/%	多态性标记数 Number of polymorphic marker	多态率 Polymorphic rate/%
单核苷酸Mononucleotide	30	29	96.67	1	3.45
二核苷酸Dinucleotide	30	28	93.33	8	28.57
三核苷酸Trinucleotide	30	28	93.33	6	21.43
四核苷酸Tetranucleotide	30	29	96.67	4	13.79
五核苷酸Tetranucleotide	30	29	96.67	7	24.14
六核苷酸Hexanucleotide	30	26	86.67	6	23.08
总计Total	180	169	93.89	32	18.93

SSR类型 SSR type	数量 Number	扩增引物数量 Number of amplified marker	扩增率 Amplification rate/%	多态性标记数 Number of polymorphic marker	多态率 Polymorphic rate/%
单核苷酸Mononucleotide	30	29	96.67	1	3.45
二核苷酸Dinucleotide	30	28	93.33	8	28.57
三核苷酸Trinucleotide	30	28	93.33	6	21.43
四核苷酸Tetranucleotide	30	29	96.67	4	13.79
五核苷酸Tetranucleotide	30	29	96.67	7	24.14
六核苷酸Hexanucleotide	30	26	86.67	6	23.08
总计Total	180	169	93.89	32	18.93

序号 Number	标记名称 Marker name	SSR基序单元核苷酸数量 Nucleotide number of SSR motif unit	正向引物 Forward primer(5'→3')	反向引物 Reverse primer(5'→3')	多态性信息含量 Polymorphism information content
1	MC03_38098	5	TTGACCCATAAATTTTGCCC	AGTCGTCGGAACACTGGTTT	0.42
2	MC05_59428	5	CAGTGCATTTCGCTCTTCAA	TTGTGCGAGTCAATTCTTGC	0.07
3	MC05_60613	5	ATCTGCCACGTTTAGGTTGG	TCATCAACACCCAGTGGAAA	0.25
4	MC06_91895	5	TTCTCTTTCTCACCTCCCGA	GTAGAAGGAGGGAGTTGGGG	0.37
5	MC06_91975	5	CACATTCACAACGCACCTTC	CCAATTTTCCTTCTGGGTTG	0.37
6	MC10_145502	5	GTGGATGCATGGAGATGATG	TCCAACATTGGGCTACTTGA	0.13
7	MC01_10177	2	TCTTCATTGCTTTTGGCCTT	TTATGTGCAGCCAAATTCCA	0.57
8	MC01_10442	2	AATGCAACCAGCTCAAATCC	GGTTAGCTTGTCGGTGGTGT	0.55
9	MC01_10775	2	GGAAATTGGGATTCGTCTGA	GCGTTCAGTTCTCGGAAAAA	0.47
10	MC03_33224	2	GGCAGAATGGGACATGAACT	AACCGAGTCAAGTACAGGGG	0.07
11	MC03_36734	2	ACCAGTTGCCCTAACTGTGG	GGCAATGATTTTGAATGCG	0.60
12	MC04_41838	2	GGTCCCCACTGTTTCCTGTA	TCAGCTGCCATATTCAGTGC	0.60
13	MC04_50530	2	TGCAATACTCTACCCCACACA	CGCCCCACCACACTAGAATA	0.70
14	MC04_52709	2	GGATTCTCCCACAGTAGGCA	CAAGGACACAGTCTCCCTCC	0.19
15	MC04_56695	2	ATTGGTGGCCTTTTGTTGTC	TGAGAAGTCAAAGTTGGGGG	0.37
16	MC05_66719	2	AACCCCTGCAAAATTCCATT	TTTCCTTGAGACGGTGATCC	0.62
17	MC05_66898	2	GCTTAATTTTCTTTTGGTCCCC	TCAAATTTGTAGGCCCTTGG	0.30
18	MC05_69589	2	AGAGACGCAAGGAAGATCCA	ATTGTGGGAGGAGCTCAATG	0.42
19	MC05_69594	2	TGTGTAGTGATGGGACAAGACTTT	TGGATAGCTTCATACCGTGTCA	0.75
20	MC06_73082	2	GCAGGCGAACATCCTACATT	CAAAACCACCTCCCTGAAGA	0.27
21	MC06_74604	2	TGGCAACCTAACCTATACTTGC	ACCAAGAGGTTATGCGTTCG	0.07
22	MC06_76026	2	CTCCTCCATCCCAATTCTCA	TCTCTTCCACACCAAATCCTC	0.07
23	MC06_82550	2	TGCTTGGTCAATTTTGTCAC	CTTTTTCACGGCGAGTCTTC	0.12
24	MC06_82800	2	TGATTGTTATGGCTGCTCCA	GCACCAGTCAATGGCTACAG	0.13
25	MC06_85886	2	TTTTCCTGTAATGCGACCGT	TCGAGTTGGTGATTGGTTGA	0.37
26	MC06_87314	2	CCAAGAAGGAAGCCAACAGA	AGGAGAAATCCCACGAAACT	0.70
27	MC06_88695	2	GGGGTGGTATTCCTGAAACA	CATTCAACCTTCAACAGGGT	0.07
28	MC06_88956	2	CCTCAATCAACAAGGGGAAA	ATGAGCACTGTCTCCAGCCT	0.47
29	MC06_89197	2	CGGTCGAAAAATCAAACCAC	TATGCCGCATGTATCGTTGT	0.13
30	MC06_91328	2	TGATCCACCTCCTTCTACGTG	GGCATGTCGTTGATCGTACC	0.52
31	MC07_98759	2	TTCACAACTGTTGCTGTGGAG	CCAAAACATTAGAGAGACCGAA	0.43
32	MC08_118245	2	TGCTTGCTCGAGGTTTCTCT	TGTTCAAATCTCCTCCCCAA	0.42
33	MC08_125161	2	GGGTTGCTCAGTTGGAGAAG	CTGCAAAACCCCATGAAAAT	0.40
34	MC09_128186	2	TTGGACAAAGCCACATGCTA	TATTTCTCGCCGACGTTAGG	0.51
35	MC10_142490	2	CCACATAACACATGGGACGA	AAATCTGAGGTCGGAGTCCA	0.35
36	MC10_145501	2	AAATCCAACTGTGCTTTGCC	CATCATCTCCATGCATCCAC	0.40
37	MC10_146038	2	GGCCTGCGACAAATAAAAGA	TCTTCACCCATCTCAAAGGG	0.55
38	MC11_155754	2	ATAGGATCACGGCGAATCAG	TCGCACTTGTTCAATTCTGC	0.13
39	MC11_156296	2	TGCAAGTTTCATTGCACCTA	CGTCTTTGTCTGGTTGCTCA	0.13
40	MC11_159185	2	CCGAATTTTTGTGGATTTCG	GGATCCATTTTGATGCCTTG	0.16
	平均Mean				0.36

Genetic diversity analysis and fingerprint construction of Dading bitter gourd germplasm resources

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标记组合 Marker combination	区分材料编号 Number of distinguished materials	区分开的材料累计数量 Cumulative number of distinguished materials
MC05_69594	19+24+26	3
MC05_69594+MC04_50530	1+7+9+10+11+12+14+15+16+17+22+23	15
MC05_69594+MC04_50530+MC06_87314	5+6+18+25	19
MC05_69594+MC04_50530+MC06_87314+MC10_146038	4+8+13+20+21	24
MC05_69594+MC04_50530+MC06_87314+MC10_146038+…	2+3?	26

编号 Number	材料名称 Material name	标记代码¹⁾ Marker code
1	东来大顶Donglai	1-1-1-1
2	1号大顶NO.1	2-2-2-2
3	3号大顶NO.3	2-2-2-2
4	翠绿三号Cuilyu NO.3	3-3-3-3
5	清蔬2号Qingshu NO.2	2-1-1-4
6	金利塔Jinlita	2-1-2-2
7	巨顶大顶Juding	4-4-3-4
8	珍珠大顶Zhenzhu	5-3-4-2
9	超绿大顶Chaolyu	2-4-3-2
10	科丰大顶Kefeng	2-1-3-2
11	利农珍珠LNzhenzhu	2-5-3-2
12	博优特804 BYT 804	4-3-1-2
13	翠绿厚肉Cuilyu HR	5-3-4-3
14	金顶3号Jinding NO.3	6-1-3-3
15	华鼎3号Huading NO.3	2-3-2-2
16	真功夫13号Zhengongfu NO.13	7-5-3-2
17	特大大顶Teda	1-3-3-2
18	青蕾1号Qinglei NO.1	7-1-3-2
19	美鼎8号Meiding NO.8	8-1-5-2
20	江门大顶Jiangmen	3-3-3-2
21	鼎胜大顶Dingsheng	5-3-4-5
22	谭边大顶Tanbian	7-3-2-2
23	广良2号Guangliang NO.2	3-1-1-4
24	江门F Jiangmen F	9-3-2-2
25	惠州F Huizhou F	7-1-6-6
26	苹果大顶Pingguo	10-3-7-2

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