广东省中山市五桂山土沉香空间遗传结构
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摘要
为了解五桂山的土沉香(Aquilaria sinensis)种群空间遗传结构,利用18个微卫星位点对143株胸高直径大于5 cm的土沉香个体基因型进行了检测。结果表明,五桂山土沉香种群的观测杂合度和期望杂合度均为0.534;近交系数为0.000 1,说明五桂山土沉香遗传多样性并不低,且种群处于随机交配状况。空间遗传结构分析结果表明,五桂山土沉香缺乏空间遗传结构,不同遗传背景的个体相互混杂,导致相邻个体遗传差异较大。这可能是五桂山土沉香种群发育过程中自疏所致,也可能是这一地区土沉香为人工种植,而非自然发育形成而造成的。
In order to understand the spatial genetic structure of Aquilaria sinensis in Wuguishan, Zhongshan City, Guangdong Province, the genotypes of 143 individuals of A. sinensis with DBH(diameter at breast height) more than 5 cm were determined by using 18 microsatellite loci. The results showed that the observed heterozygosity and expected heterozygosity of A. sinensis population were all 0.534, and inbreeding coefficient was 0.000 1, which indicated that the genetic diversity of A. sinensis in Wuguishan was not low, and the whole population was under random mating situation. The genetic landscape shapes, spatial principal components analysis(sPCA) and spatial autocorrelation analysis showed A. sinensis in Wuguishan lack spatial genetic structure. The individuals with different genetic backgrounds mixed, so that the neighbors had high genetic differences. So, it was suggested that the self thinning and artificial planting of A. sinensis could be the two possible reasons.
In order to understand the spatial genetic structure of Aquilaria sinensis in Wuguishan, Zhongshan City, Guangdong Province, the genotypes of 143 individuals of A. sinensis with DBH(diameter at breast height) more than 5 cm were determined by using 18 microsatellite loci. The results showed that the observed heterozygosity and expected heterozygosity of A. sinensis population were all 0.534, and inbreeding coefficient was 0.000 1, which indicated that the genetic diversity of A. sinensis in Wuguishan was not low, and the whole population was under random mating situation. The genetic landscape shapes, spatial principal components analysis(sPCA) and spatial autocorrelation analysis showed A. sinensis in Wuguishan lack spatial genetic structure. The individuals with different genetic backgrounds mixed, so that the neighbors had high genetic differences. So, it was suggested that the self thinning and artificial planting of A. sinensis could be the two possible reasons.
引文
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[24] HU C. Seed dispersal in Aquilaria sinensis by paper wasp[D]. Nanchang:Nanchang University, 2012:1–57.胡超.胡蜂对白木香种子传播的作用[D].南昌:南昌大学, 2012:1–57.
[25] MANOHARA T N. Wasp-mediated seed dispersal in agarwood plant(Aquilariamalaccensis),acriticallyendangeredandoverexploited species of north east India[J]. Curr Sci, 2013, 105(3):298–299.
[26] HUANG J X, LIU X K, YE Y C, et al. Leaf morphology and genetic diversity of Aquilaria sinensis in Dongguan and its surrounding areas[J]. Guangdong Agric Sci, 2014, 41(3):153–158. doi:10.3969/j.issn.1004-874X.2014.03.037.黄久香,刘宪宽,叶永昌,等.东莞与邻近地区土沉香居群的叶形态和遗传多样性分析[J].广东农业科学, 2014, 41(3):153–158. doi:10.3969/j.issn.1004-874X.2014.03.037.
[27] WRIGHT S. Evolution and the Genetics of Populations[M]. Chicago:University of Chicago Press, 1978:1–590.
[28] QICH,JINZX,LIJM.Small-scalespatialpatternsofgenetic structureinCastanopsiseyreipopulationsbasedonautocorrelation analysis in the Tiantai Mountain of Zhejiang Province[J]. Acta Ecol Sin, 2011, 31(18):5130–5137.祁彩虹,金则新,李钧敏.浙江天台山甜槠种群遗传结构的空间自相关分析[J].生态学报, 2011, 31(18):5130–5137.
[29] BERENSDG,BRAUNC,GONZáLEZ-MARTíNEZSC,etal.Fine-scale spatial genetic dynamics over the life cycle of the tropical treePrunusafricana[J].Heredity,2014,113(5):401–407.doi:10.1038/hdy.2014.40.
[30] CHUNG M Y, EPPERSON B K, CHUNG M G. Genetic structure of age classes in Camellia Japonica(Theaceae)[J]. Evolution, 2003, 57(1):62–73. doi:10.1111/j.0014-3820.2003.tb00216.x.
[31] MURRELL D J.On the emergent spatial structure of size-structured populations:When does self-thinning lead to a reduction in clustering?[J]. J Ecol, 2009, 97(2):256–266. doi:10.1111/j.1365-2745.2008.01475.x.
[32] ROBERTS D G, OTTEWELL K M, WHELAN R J, et al. Is the postdisturbance composition of a plant population determined by selection for outcrossed seedlings or by the composition of the seedbank?[J].Heredity, 2014, 112(4):409–414. doi:10.1038/hdy.2013.119.
[33] GONZáLEZ-DíAZP,JUMPAS,PERRYA,etal.Ecologyand managementhistorydrivespatialgeneticstructureinScotspine[J].For Ecol Manage, 2017, 400:68–76. doi:10.1016/j.foreco.2017.05.035.
[34] GUIDUGLI M C, NAZARENO A G, FERES J M, et al. Small but not isolated:ApopulationgeneticsurveyofthetropicaltreeCariniana estrellensis(Lecythidaceae)in a highly fragmented habitat[J]. Heredity,2016, 116(3):339–347. doi:10.1038/hdy.2015.108.
[35] VEKEMANSX,HARDYOJ.Newinsightsfromfine-scalespatial genetic structure analyses in plant populations[J]. Mol Ecol, 2004, 13(4):921–935. doi:10.1046/j.1365-294X.2004.02076.x.
[36] SCHWARTZ M K, McKELVEY K S. Why sampling scheme matters:the effect of sampling scheme on landscape genetic results[J]. Conserv Genet, 2009, 10(2):441–452. doi:10.1007/s10592-008-9622-1.
[37] ZENG L Y, XU L L, TANG S Q, et al. Effect of sampling strategy on estimationoffine-scalespatialgeneticstructureinAndrosacetapete(Primulaceae), an alpine plant endemic to Qinghai-Tibetan Plateau[J].JSystEvol,2010,48(4):257–264.doi:10.1111/j.1759-6831.2010.00084.x.
[2]MEI Q X, LI H C, WANG K Y. Certificate of origin and development conception of Aquilaria sinensis in Zhongshan[J]. Lishizhen Med Mat Med Res, 2007, 18(8):2049–2051. doi:10.3969/j.issn.1008-0805.2007.08.166.梅全喜,李汉超,汪科元.南药中山沉香的产地考证与发展构想[J].时珍国医国药,2007,18(8):2049–2051.doi:10.3969/j.issn.1008-805.2007.08.166.
[3]MEIQX,LIHC,WANGKY,etal.Medicalhistoryandorigin certificate of Aquilaria sinensis[J]. Pharm Today, 2011, 21(1):3–5.梅全喜,李汉超,汪科元,等.南药沉香的药用历史与产地考证[J].今日药学, 2011, 21(1):3–5.
[4]MEI Q X. Fragrant Medicines:Aquilaria sinensis[M]. Beijing:China Press of Traditional Chinese Medicine, 2016:1–240.梅全喜.香药:沉香[M].北京:中国中医药出版社, 2016:1–240.
[5]WANG Z F,LIAN J Y, HUANG GM,et al. Genetic groups in the commonplantspeciesCastanopsischinensisandtheirassociations with topographic habitats[J]. Oikos, 2012, 121(12):2044–2051. doi:10.1111/j.1600-0706.2012.20483.x.
[6]WANG Z F, LIAN J Y, YE W H, et al. The spatial genetic pattern of Castanopsischinensis in a large forest plot with complex topography[J]. For Ecol Manage, 2014, 318:318–325. doi:10.1016/j.foreco.2014.01.042.
[7]WANG Z F, LIAN J Y, YE W H, et al. Pollen and seed flow under differentpredominantwindsinwind-pollinatedandwind-dispersed species Engelhardia roxburghiana[J]. Tree Genet Genomes, 2016, 12(2):19. doi:10.1007/s11295-016-0973-3.
[8]JIANQC,TANZJ,LIAOHB,etal.Thegeneticdiversityof Aquilaria sinensis in Wuguishan, Zhongshan, Guangdong Province[J].Guihaia, 2018, 38(6):804–811. doi:10.11931/guihaia.gxzw201709027.蒋谦才,谭宗健,廖浩斌,等.广东省中山市五桂山土沉香(Aquilaria sinensis)遗传多样性[J].广西植物, 2018, 38(6):804–811.doi:10.11931/guihaia.gxzw201709027.
[9]PEAKALL R, SMOUSE P E. GenAlEx 6.5:Genetic analysis in Excel.Population genetic software for teaching and research:An update[J].Bioinformatics, 2012, 28(19):2537–2539. doi:10.1093/bioinformatics/bts460.
[10] ROUSSETF.GENEPOP'007:Acompletere-implementationofthe GENEPOPsoftwareforWindowsandLinux[J].MolEcolResour,2008, 8(1):103–106. doi:10.1111/j.1471-8286.2007.01931.x.
[11] WEIR B, COCKERHAM C. Estimating F-statistics for the analysis of population structure[J]. Evolution, 1984, 38(6):1358–1370. doi:10.2307/2408641.
[12] HOLM S. A simple sequentially rejective multiple test procedure[J].Scand J Statist, 1979, 6(2):65–70.
[13] MILLER M P. Alleles In Space(AIS):Computer software for the joint analysis of interindividual spatial and genetic information[J]. J Hered,2005, 96(6):722–724. doi:10.1093/jhered/esi119.
[14] JOMBART T, DEVILLARD S, DUFOUR A B, et al. Revealing cryptic spatial patterns in genetic variability by a new multivariate method[J].Heredity, 2008, 101(1):92–103. doi:10.1038/hdy.2008.34.
[15] COLE C T. Genetic variation in rare and common plants[J]. Annu Rev EcolEvol Syst,2003,34:213–237.doi:10.1146/annurev.ecolsys.34.030102.151717.
[16] NYBOMH.ComparisonofdifferentnuclearDNAmarkersfor estimating intraspecific genetic diversity in plants[J]. Mol Ecol, 2004,13(5):1143–1155. doi:10.1111/j.1365-294X.2004.02141.x.
[17] CHENG,LIUCQ,SUNWB.Pollinationandseeddispersalof Aquilariasinensis(Lour.)Gilg(Thymelaeaceae):Aneconomicplant specieswithextremelysmallpopulationsinChina[J].PlantDivers,2016, 38(5):227–232. doi:10.1016/j.pld.2016.09.006.
[18] WARDM,DICKCW,GRIBELR,etal.Toself,ornottoself:A reviewofoutcrossingandpollen-mediatedgeneflowinneotropical trees[J]. Heredity, 2005, 95(4):246–254. doi:10.1038/sj.hdy.6800712.
[19] ISMAIL S A, GHAZOUL J, RAVIKANTH G, et al. Does long-distance pollendispersalprecludeinbreedingintropicaltrees?Fragmentation geneticsofDysoxylummalabaricuminanagro-forestlandscape[J].Mol Ecol, 2012, 21(22):5484–5496. doi:10.1111/mec.12054.
[20] FINGER A, KAISER-BUNBURY C N, KETTLE C J, et al. Genetic connectivity of the moth pollinated tree Glionnetia sericea in a highly fragmented habitat[J]. PLoS One, 2014, 9(10):e111111. doi:10.1371/journal.pone.0111111.
[21] NOREEN A M E, NIISSALO M A, LUM S K Y, et al. Persistence of long-distance, insect-mediated pollen movement for a tropical canopy treespeciesinremnantforestpatchesinanurbanlandscape[J].Heredity, 2016, 117(6):472–480. doi:10.1038/hdy.2016.64.
[22] HANSON T R, BRUNSFELD S J, FINEGAN B, et al. Pollen dispersal andgeneticstructureofthetropicaltreeDipteryxpanamensisina fragmented Costa Rican landscape[J]. Mol Ecol, 2008, 17(8):2060–2073. doi:10.1111/j.1365-294X.2008.03726.x.
[23] CHáVEZ-PESQUEIRA M, SUáREZ-MONTES P, CASTILLO G, et al. Habitat fragmentation threatens wild populations of Carica papaya(Caricaceae)in a lowland rainforest[J]. Amer J Bot, 2014, 101(7):1092–1101. doi:10.3732/ajb.1400051.
[24] HU C. Seed dispersal in Aquilaria sinensis by paper wasp[D]. Nanchang:Nanchang University, 2012:1–57.胡超.胡蜂对白木香种子传播的作用[D].南昌:南昌大学, 2012:1–57.
[25] MANOHARA T N. Wasp-mediated seed dispersal in agarwood plant(Aquilariamalaccensis),acriticallyendangeredandoverexploited species of north east India[J]. Curr Sci, 2013, 105(3):298–299.
[26] HUANG J X, LIU X K, YE Y C, et al. Leaf morphology and genetic diversity of Aquilaria sinensis in Dongguan and its surrounding areas[J]. Guangdong Agric Sci, 2014, 41(3):153–158. doi:10.3969/j.issn.1004-874X.2014.03.037.黄久香,刘宪宽,叶永昌,等.东莞与邻近地区土沉香居群的叶形态和遗传多样性分析[J].广东农业科学, 2014, 41(3):153–158. doi:10.3969/j.issn.1004-874X.2014.03.037.
[27] WRIGHT S. Evolution and the Genetics of Populations[M]. Chicago:University of Chicago Press, 1978:1–590.
[28] QICH,JINZX,LIJM.Small-scalespatialpatternsofgenetic structureinCastanopsiseyreipopulationsbasedonautocorrelation analysis in the Tiantai Mountain of Zhejiang Province[J]. Acta Ecol Sin, 2011, 31(18):5130–5137.祁彩虹,金则新,李钧敏.浙江天台山甜槠种群遗传结构的空间自相关分析[J].生态学报, 2011, 31(18):5130–5137.
[29] BERENSDG,BRAUNC,GONZáLEZ-MARTíNEZSC,etal.Fine-scale spatial genetic dynamics over the life cycle of the tropical treePrunusafricana[J].Heredity,2014,113(5):401–407.doi:10.1038/hdy.2014.40.
[30] CHUNG M Y, EPPERSON B K, CHUNG M G. Genetic structure of age classes in Camellia Japonica(Theaceae)[J]. Evolution, 2003, 57(1):62–73. doi:10.1111/j.0014-3820.2003.tb00216.x.
[31] MURRELL D J.On the emergent spatial structure of size-structured populations:When does self-thinning lead to a reduction in clustering?[J]. J Ecol, 2009, 97(2):256–266. doi:10.1111/j.1365-2745.2008.01475.x.
[32] ROBERTS D G, OTTEWELL K M, WHELAN R J, et al. Is the postdisturbance composition of a plant population determined by selection for outcrossed seedlings or by the composition of the seedbank?[J].Heredity, 2014, 112(4):409–414. doi:10.1038/hdy.2013.119.
[33] GONZáLEZ-DíAZP,JUMPAS,PERRYA,etal.Ecologyand managementhistorydrivespatialgeneticstructureinScotspine[J].For Ecol Manage, 2017, 400:68–76. doi:10.1016/j.foreco.2017.05.035.
[34] GUIDUGLI M C, NAZARENO A G, FERES J M, et al. Small but not isolated:ApopulationgeneticsurveyofthetropicaltreeCariniana estrellensis(Lecythidaceae)in a highly fragmented habitat[J]. Heredity,2016, 116(3):339–347. doi:10.1038/hdy.2015.108.
[35] VEKEMANSX,HARDYOJ.Newinsightsfromfine-scalespatial genetic structure analyses in plant populations[J]. Mol Ecol, 2004, 13(4):921–935. doi:10.1046/j.1365-294X.2004.02076.x.
[36] SCHWARTZ M K, McKELVEY K S. Why sampling scheme matters:the effect of sampling scheme on landscape genetic results[J]. Conserv Genet, 2009, 10(2):441–452. doi:10.1007/s10592-008-9622-1.
[37] ZENG L Y, XU L L, TANG S Q, et al. Effect of sampling strategy on estimationoffine-scalespatialgeneticstructureinAndrosacetapete(Primulaceae), an alpine plant endemic to Qinghai-Tibetan Plateau[J].JSystEvol,2010,48(4):257–264.doi:10.1111/j.1759-6831.2010.00084.x.