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题名: 耳石形态和元素组成及其与鱼类群体识别的研究
作者: 张国华
答辩日期: 2000
导师: 曹文宣
专业: 水生生物学
授予单位: 中国科学院水生生物研究所
授予地点: 中国科学院水生生物研究所
学位: 博士
关键词: 耳石 ; 形态 ; 元素组成 ; 群体识别 ; 种类识别 ; 鱼类
摘要: 通过研究鱼类耳石的形态或元素组成,可以识别鱼类的不同群体,再现鱼类生活史,从而准确地评估鱼类资源。1997年8月-1999年5月在长江上中下游以及附属湖泊等12个区域内,共采集鱼类18种,1578号样本。研究内容包括耳石的形态、传统形态方法和傅立叶形态分析方法对鱼类种类和群体的识别、生物因素对耳石形态和元素含量的影响、耳石元素的区域差别以及这种差别在鱼类群体识别和洄游研究的应用。主要结果如下:1.研究并描述了16种鲤科鱼类微耳石、星耳石的形态。耳石具有种的鉴别特征,同一亚科不同种类的形态特征相似,亚科之间一般存在可区分的特征。因此,耳石形 态可能可以反映鲤科鱼类的系统发育关系。2.用传统形态方法对6个种类和三个区域鲫群体进行了研究。耳石形态存在体长、年龄效应。微耳石对6个种类的判别成功率为95.24%,星耳石为97.57%,微耳石和星耳石的联合判别效果为99.10%。运用耳石形态性状对三个区域鲫群体的判别成功率为微耳石70.00%,星耳石60.00%,微耳石和星耳石的联合判别效果为68.90%,这说明了鲫群体之间的耳石形态存在差异,在一定程度上可通过形态特征的比较,将鱼类群体区分开。传统形态方法是鱼类种类和群体识别的有效工具。3.用傅立叶形态分析方法对8个种类耳石形态的判别结果表明,种类之间的耳石基本轮廓存在明显的差异,这些差异可以由傅立叶前5组谐值表达出来,86%以上的个体可以通过前5组谐值正确地归入所属的种类,第5组之后的谐值对提高判别成功率是有贡献的。用微耳石、星耳石和两者联合性状对8个种类判别成功率分别为95.50%、93.95%和98.92%,傅立叶形态分析方法在种类判别效果上与传统形态方法相同。4.用傅立叶形态分析方法对七个区域鲤、三个区域鲫和三个区域草鱼耳石形态进行了研究。体长、性别、年龄对所研究群体的耳石形态存在统计意义上的影响,但是,这种影响较低,有影响的谐值一般占谐值总的数量的10%以下,并且受影响的谐值在群体之间的出现情况不具有普遍性和规律性。因此,群体之间在体长、性别、年龄上的差异不会对群体判别结果产生影响。与种类识别相比,群体识别需要更多的谐值数量来达到相似的识别成功率。除了前5组谐值.群体识别需要描述耳石形态细节的后15组谐值,后者对群体识别的重要性远大于其对种类识别研究中的作用。利用微耳石傅立叶谐值,对七个区域鲤的判别正确率为89.94%,对三个区域鲫的判别正确率为97.12%,对三个区域草鱼的判别正确率为100%;用星耳石的判别正确率,鲤为83.85%,鲫为97.30%,草鱼为100%;用微耳石和星耳石联合判别正确率,鲤为92.45%,鲫为99.04%,草鱼为100%。以上结果表明,七个区域鲤、三个区域鲫和三个区域草鱼应分属不同的群体。微耳石、星耳石的判别效果基本相同,两者联合判别效果略高于各自单独使用的效果。傅立叶形态分析方法对群体识别的能力优于传统的形态分析方法。5.分别分析了体长、年龄、性别等因素对鲤、铜鱼和草鱼耳石元素比值的影响,结果表明,这些因素仅仅影响一些区域群体个别元素的比值。因此,可以认为群体之间所存在的体长、年龄、性别上的差异不会对本研究的结果产生影响。6.通过对鲤、铜鱼和草鱼区域群体之间耳石元素比值的比较分析,发现同种鱼类不同区域群体之间普遍存在差别,这种差别可以应用于群体的识别。邻近区域群体之间耳石元素的比值差异程度较小,而间隔较远的区域群体之间元素的比值差异较大。七个区域鲤群体的判别成功率为71.60%,~(85)Rb/~(44)Ca和Sr、Mg的同位素与~(44)Ca的比值在鲤群体识别中起到了重要的作用。铜鱼六个区域的判别成功率为66.67%,Tl、Sr和Ga是铜鱼耳石元素中区域差异较大的元素。草鱼三个区域判别成功率为93.55%,Sr、Na、Mg和Ba是区分草鱼区域群体的主要元素。7.簰洲和洞庭湖鲤耳石~(23)Na/~(44)Ca、~(24)Mg/~(44)Ca、~(69)Ga/~(44)Ca、~(136)Ba/~(44)Ca、~(137)Ba/~(44)Ca、~(138)Ba/~(44)Ca的比值以及簰洲~(205)Ti/~(44)Ca与温度呈正相关关系。簰洲鲤耳石~(88)Sr/~(44)Ca和洞庭湖鲤耳石~(52)Cr/~(44)Ca与温度为负相关关系。水温与鲤耳石元素比值的相关系数较低,大部分方差不能用之解释。因此,水温对鲤耳石元素沉积的影响是比较小的。无季节差异的元素比值为~(85)Rb/~(44)Ca、~(130)Te/~(44)Ca和~(200)Hg/~(44)Ca。有季节性差异的为~(23)Na/~(44)Ca、~(24)Mg/~(44)Ca、~(52)Cr/~(44)Ca、~(69)Ga/~(44)Ca、~(88)Sr/~(44)Ca和~(138)Ba/~(44)Ca。鲤耳石的~(23)Na/~(44)Ca、~(52)Cr/~(44)Ca、~(69)Ga/~(44)Ca、~(88)Sr/~(44)Ca和~(138)Ba/~(44)Ca季节变化在簰洲与洞庭湖之间无差别,而~(24)Mg/~(44)Ca、~(85)Rb/~(44)Ca、~(130)Te/~(44)Ca和~(200)Hg/~(44)Ca在两地之间存在差异。鲤耳石元素的季节变化既不能用Radtke温度决定假说完全解释,也不能用Kalish生理决定假说解释。淡水鱼类耳石元素的季节变化可能更多地受到水域环境中元素含量和饵料生物季节变化的影响。8.通过对中华鲟成鱼胸鳍骨外表(淡水生活)和截面(海洋生活),以及淡水和河口的幼鱼胸鳍骨外表所含元素的比较分析,寻找可以指示中华鲟不同生活区域的“指示元素”。任一区域的~(23)Na/~(44)Ca、~(138)Ba/~(44)Ca与其它区域都存在统计上的显著性差别,淡水生活阶段的~(138)Ba/~(44)Ca高,海洋生活阶段的~(138)Ba/~(44)Ca低,~(23)Na/~(44)Ca与~(138)Ba/~(44)Ca相反,呈淡水低、海洋高的格局,在中华鲟成鱼、幼鱼中,此两种元素比值都表现出此同样的现象。这说明中华鲟从一个环境洄游到另一个环境后(海洋→淡水,淡水→海洋),胸鳍骨中的~(23)Na/~(44)Ca、~(138)Ba/~(44)Ca会很快发生变化,~(23)Na/~(44)Ca、~(138)Ba/~(44)Ca具有快速反映环境变化的特点,可以作为中华鲟在成淡水水域之间洄游的“指示元素”。这种“指示元素”的应用将有助于揭示中华鲟的生活史如初次性成熟年龄、繁殖周期甚至其周年活动规律等。成鱼~(88)Sr/~(44)Ca比值显著高于幼鱼,说明了~(88)Sr/~(44)Ca比值的高低是中华鲟在海洋或淡水生活的识别特征;成鱼海洋生活阶段与成鱼淡水生活阶段~(88)Sr/~(44)Ca无显著性差异,幼鱼与幼鱼之间~(88)Sr/~(44)Ca虽有差异,但变化幅度较小,说明中华鲟从一个环境进入另一个环境后(海洋→淡水,淡水→海洋),~(88)Sr/~(44)Ca不会很快发生变化,即存在时滞现象。这一现象与以往对其它鱼类的研究是不一样的。不同元素对环境变化的反应存在差别现象的发现将促进国内外对“快速反应”元素的沉积规律展开研究。9.“元素指纹”技术和傅立叶图形分析方法均首次用于我国鱼类学的研究,其中运用傅立叶图形分析方法进行鱼类种类的识别在国内外当属首例。研究结果表明,这两种识别方法对种类、群体都有较高的判别成功率,其中“元素指纹”技术是研究鱼类洄游的很好的方法,可以解决用传统方法所不能解决的问题。相信随着分析仪器灵敏度的不断提高,该研究具有广阔的应用前景。与传统的形态分析方法比较,傅立叶图形分析方法具有对形态的变化更为敏感、数据分析的可软件化等优势,可以解决鱼类生态、鱼类分类研究中的许多实际问题。
英文摘要: The total number of 18 species and 1578 specimen were collected at 12 locations along the Changjiang River and in the adjacent lakes from August, 1997 to May, 1999. The dissertation described otolith morphology, compared the efficiency of two techniques (traditional otolith morphometrics and Fourier shape analysis) in species and stock discrimination, examined the effects of biotic factors on otolith shape and elemental composition, analyzed regional differences of elemental ratios with the application in stock discrimination and migratory history. The main results are as follows: 1. The otoliths (lapilli and asterisci) of sixteen cyprinid species were described. All species could be distinguished by their otoliths, thus making otoliths a powerful taxonomic tool. There existed distinguishable otolith characteristies among most subfamilies and otolith shape affinities within the subfamilies. It suggested that otolith morphology might reveal phylogenetic relationships between species, genera and subfamilies. 2. Six species and three crucian carp stocks were studied by using traditional otolith morphometrics. The effects of body length or age on otolith shape were significant while no bilateral position effects were found. Accurate classification of six species was 95.24% by lapilli, 97.57% by asterisci and 99.10% by combined variables of lapilli and asterisci. Classification accuracy of three crucian carp stocks was 70.00% by lapilli, 60.00% by asterisci and 68.90% by combined variables of lapilli and asterisci. It suggested that there were differences of otolith shape between the crucian carp stocks. Traditional otolith morphometrics is an effective tool for species and stock identity. 3. Otolith outline was digitized with an image processor and used to calculate a complex Fourier transform of the close contour. Comparison of otolith shape variables of eight cyprinid species showed that there were significant differences of otolith outlines among species that could be expressed by the first five Fourier harmonic coefficients. More than 86% individuals could be accurately assigned to the right species by using some of the first five Fourier harmonic coefficients, while the 6th-20th Fourier harmonic coefficients were useful for improving classification success. Correct classification rates were 95.50%, 93.95% and 98.92% using Fourier harmonic coefficients of lapilli, asterisci and the combination of these two otolith types respectively. Our results suggested that Fourier shape analysis is also an effective tool for species discrimination. 4. Common carp from seven sites, crucian carp and grass carp from three sites were obtained for stock discrimination with Fourier shape analysis. Otolith shape differed among body length, sex and year-classes, but these effects were minor because the variables with significant differences usually were less than 10% of the total variables, and any of these kinds of variables did not show the same trend at all sites. Body size, sex and year-classes were not the main contributions to otolith shape variations among fish stocks. Compared with species discrimination, stock discrimination required more Fourier harmonic coefficients to reach the similar classification success. Besides the first five harmonics, stock discrimination required the 6th-20th harmonics that described the roughness of the otoliths. The 6th-20th harmonics played much important role in stock discrimination as they did in species discrimination. Correct classification was 89.94% for common carp from seven sites, 97.12% for crucian carp from three sites and 100% for grass carp from three sites by using lapillar harmonics. The asteriscal otoliths appeared as good as lapillar ones in stock discrimination as to classify 83.85% of common carp, 97.30% of crucian carp and 100% of grass carp into the right sites. The variables combined with lapillar and asteriscal otoliths provided slightly better stock discrimination than did the lapillar or asteriscal otoliths alone, and the classification success was 92.45% for common carp, 99.04% for crucian carp and 100% for grass carp respectively. These results suggested that common carp, crucian carp and grass carp from different sites had the different stock origins. It also indicated that the Fourier shape analysis was superior to the traditional morphometrics in stock discrimination. 5. Elemental ratios in the lapillar otoliths of common carp from seven sites, copper carp from six sites and grass carp from three sites were measured by LA-ICP-MS. The effects of the factors such as length, age and sex on elemental ratios of three species were evaluated. The results showed that these factors affected only a few of elements in some of sites, not all sites, so that the differences of length, age and sex among the stocks might have no significant impacts on the discrimination results.6. Through the comparison of element: ~(44)Ca ratios among stocks of common carp, copper carp and grass carp, the differences of elemental ratios between stocks of the same species was ubiquitous and could be used in stock discrimination. The differences in elemental ratios between adjacent regions were of smaller magnitude than those having longer distance s. The classification success was 71.60% for common carp from seven sites while Rb, Sr and Mg: ~(44)Ca ratios were the most important in the discrimination. Tl, Sr and Ga greatly contributed to the 66.67% correct classification of copper carp from six sites. Grass carp from three sites had the highest classification success, 93.55% , and Sr, Na, Mg and Ba were the most important elements to identify grass carp stocks. 7. Correlation analysis showed that ~(23)Na/~(44)Ca, ~(24)Mg/~(44)Ca, ~(69)Ga/~(44)Ca, ~(136)Ba/~(44)Ca, ~(137)Ba/~(44)Ca, ~(138)Ba/~(44)Ca of common carp otoliths both in Paizhou and in Dongting Lake as well as ~(205)Tl/~(44)Ca in Paizhou were positively correlated with water temperature, while ~(88)Sr/~(44)Ca in Paizhou and ~(52)Cr/~(44)Ca in Dongting Lake had negative relationships with water temperature. These relationships could not explain the most of the variances for any of the elemental ratios so that the effects of water temperature on the incorporation of elements in common carp otoliths was slight.There were no seasonal variations for ~(85)Rb/~(44)Ca,~(130)Te/~(44)Ca and ~(200)Hg/~(44)Ca ratios in common carp otoliths, while there were significant differences among bimonthly samples for ~(23)Na/~(44)Ca, ~(24)Mg/~(44)Ca, ~(52)Cr/~(44)Ca, ~(69)Ga/~(44)Ca, ~(88)Sr/~(44)Ca、and ~(138)Ba/~(44)Ca data. The seasonal cycles of ~(23)Na/~(44)Ca, ~(52)Cr/~(44)Ca, ~(69)Ga/~(44)Ca, ~(88)Sr/~(44)Ca, and ~(138)Ba/~(44)Ca, in Paizhou were similar to those in Dongting Lake, but ~(24)Mg/~(44)Ca, ~(85)Rb/~(44)Ca, ~(130)Te/~(44)Ca and ~(200)Hg/~(44)Ca in Paizhou displayed different seasonal cycles from those in Dongting Lake. Neither temperature-determination hypothesis nor seasonal physiological influences (reproductive cycles) could fully explain the seasonal changes of elemental ratios in common carp otoliths. Seasonal variations of otolith composition of freshwater fish may be mainly related to the seasonal changes of elemental concentrations in environments and food abundance. 8. The possibility of the utilization of elemental ratios as indicators of life history in Chinese sturgeon was investigated. ~(23)Na/~(44)Ca and ~(138)Ba/~(44)Ca in the pectoral fin rays of Chinese sturgeon were significantly different between locations. ~(138)Ba/~(44)Ca ratios had high levels in freshwaters and lower levels in the ocean, and the pattern of ~(23)Na/~(44)Ca ratios, in contrast to that of ~(138)Ba/~(44)Ca ratios, was with lower values in freshwater and high values in the ocean. This phenomenon was convincing in either juveniles or adults, which suggested that ~(23)Na/~(44)Ca and ~(138)Ba/~(44)Ca ratios changed quickly when fish moved from the ocean into the Changjiang River or vice versa. This result indicated that ~(23)Na/~(44)Ca and ~(138)Ba/~(44)Ca could response to the changes of environments and might be useful tracers of migration between freshwaters and the ocean. Significant differences between adults and juveniles were found for ~(88)Sr/~(44)Ca ratios, indicating that the values of ~(88)Sr/~(44)Ca were indicative of freshwater or oceanic residency for Chinese sturgeon. Significant differences between freshwater and oceanic life stage were not found for ~(88)Sr/~(44)Ca ratios of adults, whereas the ratios of juveniles decreased slightly from freshwater to the estuary, indicating that ~(88)Sr/~(44)Ca ratios in pectoral fin ray might not have responded immediately to the environmental changes when Chinese sturgeon migrated between the ocean and the Changjiang River. It is a lag period for ~(88)Sr/~(44)Ca ratios to respond to the environmental changes that is not found for other species in previous studies. 9. Our analysis of otolith shape is the first attempt to use Fourier analysis to discriminate the fish species in China and abroad. High correct classification rates were achieved with these two methods, and the "elemental fingerprint" technique is a valuable tool in solving the problems of fish ecology and behavior studies that may not be solved with other methods. Compared with traditional otolith morphometrics, Fourier shape analysis is more sensitive to otolith shape changes and shows promise as a powerful aid in the management of fisheries.
语种: 中文
内容类型: 学位论文
URI标识: http://ir.ihb.ac.cn/handle/342005/12544
Appears in Collections:中科院水生所知识产出(2009年前)_学位论文

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Recommended Citation:
耳石形态和元素组成及其与鱼类群体识别的研究.张国华[d].中国科学院水生生物研究所,2000.20-25
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