海洋纤毛虫的多样性研究:裸口类、下口类、吸管类、旋唇类
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摘要
由于高度的细胞结构分化、表型水平上的形态多样性、独特的核双态性以及(有性的)接合生殖方式、及在微食物网中的枢纽地位等因素,纤毛虫原生动物一直是细胞学、真核生物遗传学、生态学等基础生物学领域中重要的模型动物或研究对象。
     在国家自然科学基金项目(30430090,40676076)及中-英国际合作达尔文启动计划(14-015)的资助下,作者于2006-2009年期间,对中国黄渤海和南海近岸不同生境的自由生纤毛虫的区系开展了调查。结合活体观察及蛋白银、氨银法等现代染色方法对所涉纤毛虫的活体形态、纤毛图式或银线系及统计学特征进行了详细的研究和报道。共发现、分离和鉴定自由生海洋纤毛虫46种,分别隶属于9个目(前口目、刺钩目、合膜目、篮管目、管口目、吸管目、异毛目、丽克虫目、腹毛目)。根据形态学所给出的重要指征建立了若干新的分类阶元,包括2个新属、11个新种和4个新组合。
     主要成果和创新点包括:
     1)榴弹虫科,因许多重要分类特征(口器数目、排布,口区结构等)在活体观察中难以了解,造成该类群的分类学长期存在混乱和错误。本工作基于活体观察及细胞学染色制片,首次揭示了1新属(偏榴弹虫属)、4新种(东方诺兰德虫,中华诺兰德虫,巨大偏榴弹虫,相似巨大偏榴弹虫)及1新组合(相似瓦榴弹虫)的形态学和纤毛图式特征,并首次给出了迄今不明的纺锤披巾虫的口器结构。其中,新属(偏榴弹虫)特征为:具棘刺,盔板8环;盔板具肾形单窗口,脊靠近波浪状左边缘,且呈翼状,盔板右边缘的齿状结构与窗口交替排列;口器3个。
     2)吸管亚纲是纤毛门中高度特化的类群,大部分阶元尤其是海洋类群近半个世纪以来一直无人问津。本文借助银染技术及分子信息完成了对14种的研究并建立1新属(拟粘管虫属)与1新种(半球拟粘管虫):生芽斑吸管虫,冠状斑吸管虫,球形斑吸管虫,柱状斑吸管虫,结节壳吸管虫,扁平壳吸管虫,粗壮壳吸管虫,具边似壳吸管虫,宽口似壳吸管虫,钟状贝吸管虫,锯齿杯吸管虫,小甲吸管虫,长号甲吸管虫,灰色曲壳吸管虫。拟粘管虫新属的特征为:虫体半球形,无柄、无鞘,通过基板直接附着在基质上;细胞明显分成致密的内质和透明的外质,后者呈凝胶状。吸管触手状,不成簇,散布于细胞表面。
     3)异毛目隶属于分类地位最高等的多膜纲,虫体通常巨大,常见于各类水体,其研究史几乎贯穿了纤毛虫的研究史。因其各种类间活体特征的重叠性,在300多年的历史中积累了大量混乱和错误。本工作基于活体观察及详尽的纤毛图式资料,激活长期存在争议的突喇叭虫属且给出新定义,并对其模式种(耳状突喇叭虫)与绿色爽口虫进行了混乱清理、重认证及缺失资料的补足,同时,对8个已知种(蓝菌丽克虫,简单瓶囊虫,延长类瓶囊虫,佛氏环须虫,盐蚕豆虫,小突口虫,叶状突口虫,弯曲突口虫)进行了必要性重描述及显微照片的提供。突喇叭虫新激活属的特征为:虫体较大并具一定伸缩性,自由游泳;深而阔大的口前庭,前庭腹面边缘有一裂口;口前庭动基列仅分布在口前庭内壁上,口区底部没有毛基粒或动基列;纤毛图示与喇叭虫相似:具腹面缝合线和体动基列条带反差区;口围带几乎闭合,在腹面形成一楔形缺口;口侧膜长而明显。
     4)合膜目种类相对较少,虽仅有7属,但大部分种类缺少纤毛图式及明确的分类标准,各属种间分类与系统地位长期不明。借助银染技术及分子手段,对该类群前人遗留的混乱做了清理和厘定,建立3个新组合(椭圆带膜虫,贪食带膜虫,长形带膜虫),给出该目种类最多属(带膜虫属)的新定义:虫体肾形或卵圆形;合膜粗壮,口区左右两侧均有分布,延伸到左侧背面;大核椭球形或长棒状;伸缩泡通常多个,位置不确定。
     5)对管口目类群做了补足性研究,建立3个新种(热带管口虫,喙突宽管虫,似梳状偏体虫),对国际间长期存疑,分类学资料不详的3个种做了纤毛图式资料的建立和重描述,分别给出新定义、建立新模标本(北欧林奇虫,少齿篷管虫,寄生下管虫)。
     6)前口目隶属裸口亚纲,分类地位原始,彼此间亲缘关系不明或存严重争议。本工作在详尽活体观察基础上,借助蛋白银等银染技术对该类群进行了初步探讨。建立1个新种(拟纹中圆虫),并首次给出迄今不明的优美壶形虫的纤毛图式和详细的口区结构,对其分类地位的确定提供了重要证据,同时对已知种盐扁体虫进行了补足性重描述以及统计数据的提订。
     7)同时,本工作对篮管目与腹毛目2个新种进行了现代方法的研究,包括活体观察、蛋白银显微照片、纤毛图式及分子信息(红色篮口虫,多核拟双棘虫),对长期分类地位不明的盾片异列虫在纤毛图式水平上进行了清理、重认证以及缺失资料的补足,并分别给出了新定义。
Ciliated protozoa is the most complex and highly differentiated among unicellularorganisms,of which some groups are closely related to occurrence of red-tide,bait-animalculture in aquaculture,disease of animal,water quality purification in polysaprobicenvironment,material cycle and energy flow in“micro-food-loop”.
     From 2006 to 2009,faunistic and taxonomic studies on free living ciliates mainly fromcoastal waters off Qingdao and Guangzhou (including sand,mollusks-culturing and-breedingwaters,tidal zone,mouth of the Pearl River,mangroves near Shenzhen) have been carried out.The morphology and infraciliature of 46 species representing 9 orders (Prostomatida,Haptorida,Synhymeniida,Nassulida,Cyrophorida,Suctorida,Heterotrichida,Licophorida,Hypotrichida),23 families and 33 genera were investigated using living observation and silverimpregnation methods.Main achievements are summarized as follows:
     Two new genera were established,Apocoleps nov.gen.and Paramucophrya nov.gen..
     Diagnosis of Apocoleps nov.gen.:Colepidae with spiny armour composed of eight tiers;plate with reniform uni-windows,ridge near left plate margin serrated and wing-like,teeth ofright plate margin arranged alternately with windows;three adoral organelles.
     Diagnosis of Paramucophrya nov gen.:body hemispherical,composed of inner and outerplasma,the outer of which is a transparent mucous coat;without stalk or lorica,attached tosubstrate;tentacles sparsely scattered.
     The complicated genus Condylostentor Jankowski,1978 which has never been studiedusing modern methods since it is originally established,is reactivated and restudied basedon investigation on the type species of this genus.
     Improved diagnosis of Condylostentor:Slightly contractile,free-swimming Stentoridae;vestibular cavity deep and prominent,apical border with conspicuous ventral groove;ciliaturein Stentor-like pattern,i.e.ventral suture and contrast zone of somatic kineties present;adoralzone almost closed,interrupted on ventral side by a deep cleft;vestibulum kineties andparoral membrane present.
     Eleven new species were established based on living morphology and infraciliature:Nolandia orientalis Chen et al.,2009;Nolandia sinica Chen et al.,2009;Apocoleps magnusChen et al.,2009;Apocoleps paramagnus Chen et al.,2009;Metacystis parastriata nov.spec.;Nassula rubra nov.spec.;Cyrtophoron tropic nov.spec.;Aegyriana rostellum nov.spec.; Dysteria pseudopectinata nov.spec.;Paramucophrya semisphaerica Chen et al.,2005;Parabirojimia multinucleata Chen et al.,2009.
     Four new combinations were suggested and their infraciliature was reveal for thefirst time which has never been studied using modern staining methods:Pinacocolepssimilis (Kahl,1933) Chen et al.,2009;Zosterodasys oblonga (Maupas,1883) nov.comb.;Zosterodasys vorax (Stokes,1887) nov.comb.;Zosterodasys elongata (Kahl,1931) nov.comb.
     Infraciliature of 8 species was revealed for the first time and their neotypes havebeen deposited.These included:Tiarinafusus (Clapar(?)de & Lachmann,1857) Bergh,1881;Lagynus elegans (Engelmann,1862) Quennerstedt,1867;Lynchella nordica Jankowski,1968;Chlamydonys paucidentatus Deroux,1976;Hypocoma parasitica Gruber,1884;Condylostentor auriculatus (Kahl,1932) Jankowski,1978;Climacostomum virens (Ehrenberg,1933) Kahl,1932;Anteholosticha scutellum (Cohn,1866) Berger,2003.
     Fifteen species of suctorid ciliates were recorded and described for the first time inChina:Ephelota gemmipara (Hertwig,1876) Bütschli,1889;E.mammillata Dons,1915;E.crustaceorum Hailer,1880;E.truncata Fraipont,1878;Acineta tuberosa Ehrenberg,1833;A.compressa Clapar(?)de & Lachmann,1859;A.foetida Maupas,1881;Paracineta limbata(Maupas,1881) Collin,1912;P.patula (Clapar(?)de & Lachmann,1861) Collin,1911;Conchacineta complatana (Gruber,1884) Curds,1985;Actinocyathula crenata (Fraipont,1878) Curds,1987;Loricophrya parva (Schulz,1932) Matthes,1956;L.tuba (Zelinka,1914)Curds,1987;Paramucophrya semisphaerica Chen et al.,2005;Flectacineta livadiana(Mereschkowsky,1881)Jankowski,1978.
     Nine species were redescribed based on characteristics of living feature andinfraciliature from the Qingdao and Guangzhou populations:Placus salinus Dietz,1964;Licnophora lyngbycola Faur(?)-Fremiet,1937;Folliculina simplex (Dons,1917) Song et al.,2003;Folliculinopsisproducta (Wright,1859) Faur(?)-Fremiet,1936;Peritromusfaurei Kahl,1932;Fabrea salina Henneguy,1890;Condylostoma minutum Bullington,1940;Condylostoma spatiosum Ozaki & Yagiu in Yagiu,1944;Condylostoma curva Burkovsky,1970.
引文
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