我国北方岩溶泉域刚毛藻的系统发育及形态学研究（英文）
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摘要
脆弱刚毛藻(Cladophora fracta)是一种大型丝状绿藻,生境分布广泛。然而,对于岩溶泉域分布的刚毛藻研究较少,它们的遗传多样性、生物地理亲缘性和生理特性都有待于深入研究。该研究对我国北方地区五个典型岩溶泉域的50个脆弱刚毛藻样本进行了形态学和分子系统学描述。主要研究目标:(1)对我国北方地区五个典型岩溶泉的刚毛藻生境进行描述;(2)根据形态学特征和分子序列对藻体进行鉴定;(3)探究生境对藻体生理特性的影响。结果表明:基于SSU和LSU序列的结果,发现所分析的50株刚毛藻个体为同一种,同时还发现了13个不同的核糖体基因型。基于SSU和LSU的系统发育树,刚毛藻属均未能形成单系分支,分布在三个不同的分支上。13个样本基因型在SSU和LSU树中的位置相似,与Cladophora vagabunda有很高的序列同源性,但是形态特征却差异很大。从显微结构结果来看,五个岩溶泉域采集到的刚毛藻在细胞直径上无显著差异,藻体的形态特征与脆弱刚毛藻相一致。但是,岩溶泉域采集的藻体细胞直径比文献报道中在湖泊和河流中采集的脆弱刚毛藻直径要大。另外,仅在两个地点(XA和ST)采集的标本中发现有假根状分枝。因此,基于形态学和分子序列的结果,将这五个泉域的刚毛藻鉴定为脆弱刚毛藻(Cladophora fracta)。
Cladophora fracta,a filamentous green macroalgal epiphyte on rhodoliths,is described from five karst springs in North China. Although Cladophora species frequently appear in karst system,their genetic diversities,biogeographical affinities and physiological properties have not been well investigated in these environments. The specific objectives of this study were as follows:( 1) Describe the habitat of the cladophora-like algae form the five karst springs;( 2) Identify the thallus to species level based on a combination of morphological characteristics and molecular sequence;( 3) Explore the morphological influence of habitat. To elucidate the biogeographical patterns in Cladophora,both morphological and molecular evidence were compared of Cladophora specimens across five study sites. Analyses of partial small subunit( SSU) and large subunit( LSU) genes revealed that the studied 50 Cladophora specimens were genetically identical species and a total of thirteen ribotypes were detected. The molecular sequencing results indicated that the examined species was highly homologous with C. vagabunda,though they shared few morphological features. The genus did not form a monophyletic clade but in three different clades both in SSU and LSU trees. The microscopic structure was more consistent with that of C. fracta. The Cladophora from the five karst springs did not show significant variation in cell dimensions. However,the species exhibited larger cell diameters than those reported from lakes. In addition,the rhizoidlike branches are only observed in two locations( XA and ST). Considering the morphological characteristics,we therefore hold our species as C. fracta.
Cladophora fracta,a filamentous green macroalgal epiphyte on rhodoliths,is described from five karst springs in North China. Although Cladophora species frequently appear in karst system,their genetic diversities,biogeographical affinities and physiological properties have not been well investigated in these environments. The specific objectives of this study were as follows:( 1) Describe the habitat of the cladophora-like algae form the five karst springs;( 2) Identify the thallus to species level based on a combination of morphological characteristics and molecular sequence;( 3) Explore the morphological influence of habitat. To elucidate the biogeographical patterns in Cladophora,both morphological and molecular evidence were compared of Cladophora specimens across five study sites. Analyses of partial small subunit( SSU) and large subunit( LSU) genes revealed that the studied 50 Cladophora specimens were genetically identical species and a total of thirteen ribotypes were detected. The molecular sequencing results indicated that the examined species was highly homologous with C. vagabunda,though they shared few morphological features. The genus did not form a monophyletic clade but in three different clades both in SSU and LSU trees. The microscopic structure was more consistent with that of C. fracta. The Cladophora from the five karst springs did not show significant variation in cell dimensions. However,the species exhibited larger cell diameters than those reported from lakes. In addition,the rhizoidlike branches are only observed in two locations( XA and ST). Considering the morphological characteristics,we therefore hold our species as C. fracta.
引文
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POSADA D,2008.jModelTest:Phylogenetic Model Averaging[J].Mol Biol Evol,25(7):1253-1256.
RONQUIST F,HUELSENBECK JP,2003.MRBAYES 3:Bayesian phylogenetic inference under mixed models[J].Bioinformatics,19(12):1572-1574.
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SINHA JP,1968.Cytotaxonomical studies on Cladophora glomerata,four freshwater forms[J].Int J Cytol,32(3-4):507-518.
TENG LH,2011.Study on morphology and molecular phylogeny of Cladophorales(Chlorophyta)along China sea coast,with Its DNA Barcoding Based on ITS and 18S rDNA Sequences[D].Beijing:Master Dissertation,Graduate University of Chinese Academy of Sciences:17.
THOMPSON JD,GIBSON TJ,PLEWNIAK F,et al.,1997.The ClustalX windows interface:flexible strategies for multiple sequence alignment aided by quality analysis tools[J].Nucleic Acids Res,25(25):4876-4882.
VAN DEN HOEK C,CHIHARA M,2000.A taxonomic revision of the marine species of Cladophora(Chlorophyta)along the coast of Japan and the Russian Far-East.National Science Museum Monographs[M].Tokyo:National Science Museum,19:242.
WHITTON BA,1970.Biology of Cladophora in freshwaters[J].Water Res,4(7):457-476.
XIA X,XIE Z,2001.DAMBE:Software package for data analysis in molecular biology and evolution[J].J Hered,30(7):1720-1728.
YOSHII Y,HANYUDA T,WAKANA I,et al.,2004.Carotenoid compositions of the Cladophora balls(Aegagropila linnaei)and some members of the Cladophorales(Ulvophyceae,Chlorophyta):their taxonomic and evolutionary implication[J].J Phycol,40(6):1170-1177.
FELSENSTEIN J,1985.Confidence limits on phylogenies:An approach using the bootstrap[J].Evolution,39(4):783-791.
GESTINARI LMDS,PEREIRA SMB,YONESHIGUE-VALEN-TIN Y,2010.Distribution of Cladophora species(Cladophorales,Chlorophyta)along the Brazilian Coast[J].Phytotaxa,14(3):22-42.
GUINDON S,GASCUEL O,2003.A simple,fast and accurate algorithm to estimate large phylogenies by maximum likelihood[J].Systematic Biol,52(5):696-704.
HANYUDA T,WAKANA I,ARAI S,et al.,2002.Phylogenetic relationships within Cladophorales(Ulvophyceae,Chlorophyta)inferred from 18S rRNA gene sequences,with special reference to Aegagropila linnaei[J].J Phycol,38(3):564-571.
HAYAKAWA Y,OGAWA T,YOSHIKAWA S,et al.,2012.Genetic and ecophysiological diversity of Cladophora(Cladophorales,Ulvophyceae)in various salinity regimes[J].Phycol Res,60(2):86-97.
HAN XR,LU RA,LI QS,et al.,1993.Karst systemresearch on large karst springs in Shanxi[M].Beijing:Geological Publishing House.[韩行瑞,鲁荣安,李庆松,等,1993.岩溶水系统--山西岩溶大泉研究[M].北京:地质出版社.]
HU HJ,WEI YX,2006.The freshwater algae of China-systematics,taxonomy and ecology[M].Beijing:Science Press:770.[胡鸿钧,魏印心,2006.中国淡水藻类:系统,分类及生态[M].北京:科学出版社:770.]
HUELSENBECK JP,BULL JJ,CUNNINGHAM CW,1996.Combining data in phylogenetic analysis[J].Trends Ecol Evol,11(4):152-158.
ICHIHARA K,SHIMADA S,MIYAJI K,2013.Systematics of Rhizoclonium-like algae(Cladophorales,Chlorophyta)from Japanese brackish waters,based on molecular phylogenetic and morphological analyses[J].Phycologia,52(5):398-410.
LELIAERT F,BOEDEKER C,PENA V,et al.,2009.Cladophora rhodolithicola sp.nov.(Cladophorales,Chlorophyta),a diminutive species from European maerl beds[J].Eur J Phycol,44(2):55-169.
LELIAERT F,ROUSSEAU F,DE REVIERS B,et al.,2003.Phylogeny of the Cladophorophyceae(Chlorophyta)inferred from partial LSU rRNA gene sequences:is the recognition of a separate order Siphonocladales justified?[J].Eur J Phycol,38(3):233-246.
LELIAERT F,DECLERCK O,VERBRUGGEN H,et al.,2007.Molecular phylogeny of the Siphonocladales(Chlorophyta:Chlorophyceae)[J].Mol Phylogenet Evol,44(3):1237-1256.
NIENHUIS PH,1975.Biosystematics and ecology of Rhizoclonium riparium(Roth)Harvey(Chlorophyceae:Cladophorales)in the estuarine area of the rivers Rhine,Meuse,and Scheldt[M].Bronder Offset B.Y.,Rotterdam:240.
POSADA D,2008.jModelTest:Phylogenetic Model Averaging[J].Mol Biol Evol,25(7):1253-1256.
RONQUIST F,HUELSENBECK JP,2003.MRBAYES 3:Bayesian phylogenetic inference under mixed models[J].Bioinformatics,19(12):1572-1574.
RUSSELL DJ,BALAZS GH,2000.Identification manual for dietary vegetation of the Hawaiian green turtle,Chelonia mydas[M].NOAA TM-NMFS-SWFSC-294:49.
SINHA JP,1968.Cytotaxonomical studies on Cladophora glomerata,four freshwater forms[J].Int J Cytol,32(3-4):507-518.
TENG LH,2011.Study on morphology and molecular phylogeny of Cladophorales(Chlorophyta)along China sea coast,with Its DNA Barcoding Based on ITS and 18S rDNA Sequences[D].Beijing:Master Dissertation,Graduate University of Chinese Academy of Sciences:17.
THOMPSON JD,GIBSON TJ,PLEWNIAK F,et al.,1997.The ClustalX windows interface:flexible strategies for multiple sequence alignment aided by quality analysis tools[J].Nucleic Acids Res,25(25):4876-4882.
VAN DEN HOEK C,CHIHARA M,2000.A taxonomic revision of the marine species of Cladophora(Chlorophyta)along the coast of Japan and the Russian Far-East.National Science Museum Monographs[M].Tokyo:National Science Museum,19:242.
WHITTON BA,1970.Biology of Cladophora in freshwaters[J].Water Res,4(7):457-476.
XIA X,XIE Z,2001.DAMBE:Software package for data analysis in molecular biology and evolution[J].J Hered,30(7):1720-1728.
YOSHII Y,HANYUDA T,WAKANA I,et al.,2004.Carotenoid compositions of the Cladophora balls(Aegagropila linnaei)and some members of the Cladophorales(Ulvophyceae,Chlorophyta):their taxonomic and evolutionary implication[J].J Phycol,40(6):1170-1177.
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