POPULATION DYNAMICS OF DIPLOSTOMUM SPATHACEUM AND ITS PREFERENCE IN INFECTING GYMNOCYPRIS SELINCUOENSIS FROM SILING LAKE, TIBET AUTONOMOUS REGION, CHINA
-
摘要: 文章通过种群动态研究匙形双穴吸虫感染在时空上的变化, 并研究该虫感染是否存在对宿主性别、左右眼及眼球不同部位的寄生偏好性, 摸清匙形双穴吸虫在色林错裸鲤中的感染情况, 分析种群消长原因, 探究其生活史策略。跨年按不同季节采捕色林错裸鲤, 记录全长、体重和性别, 采集并统计匙形双穴吸虫囊蚴数量, 计算不同时空下的感染率和平均丰度, 通过独立样本非参数检验, 判断不同性别宿主、左右眼及眼球不同部位的感染数量是否存在显著差异性, 以检验匙形双穴吸虫是否存在寄生偏好。共剖检色林错裸鲤165尾[全长28.7—49.5 cm, 平均全长(37.9±4.0) cm, 体重196.9—827.2 g, 平均体重(473.3±127.9) g, 包括雌性82尾, 雄性83尾], 共检出匙形双穴吸虫515只, 最大寄生量为32只/尾。在时间上, 匙形双穴吸虫感染率和平均丰度在2020年夏季最高, 秋季和春季呈下降趋势, 2021年夏季感染率下降, 平均丰度却呈上升趋势, 2021年秋季与2020年秋季感染率与平均丰度较为接近。在空间上, 根据色林错裸鲤全长范围, 以5 cm为分组间隔, 将其分为5个全长组, 感染率和平均丰度在25 cm≤TL<30 cm全长组最低, 在30 cm≤TL<35 cm、35 cm≤TL<40 cm和40 cm≤TL<45 cm全长组均较为接近, 在45 cm≤TL<50 cm全长组出现了较大的上升。匙形双穴吸虫对色林错裸鲤的感染, 在不同性别的宿主中不存在偏好, 对宿主的左右眼也不存在偏好性, 但在晶状体和玻璃体中存在明显的偏好性, 更多寄生于晶状体中。匙形双穴吸虫种群消长的季节动态, 与水温等环境因素、候鸟的迁徙时间和螺类的存在息息相关。随着鱼类宿主全长的增长, 眼睛体积增加, 可容纳数量更多的复口吸虫, 长期积累性接触也使感染数目更多。在45 cm≤TL<50 cm全长组出现了较大的上升, 可能是因为更大的体型代表着更大的表面积, 不仅可受到更多尾蚴的入侵, 也在一定程度上有利于色林错裸鲤在不断增加的感染水平下保持生存, 从而积累更多的匙形双穴吸虫。匙形双穴吸虫感染色林错裸鲤的寄生偏好, 是适应环境所表现出的一种生活史策略, 有利于种群的传播和繁衍。Abstract: In this paper, the population dynamics of the parasite Diplostomum spathaceum is studied to understand infections in the hosts Gymnocypris selincuoensis from Siling Lake, including seasonal and group differences in infections and reasons for changes in the population size, identify the parasite’s preferences for male/female hosts, left/right eyes of hosts, or specific parts of hosts’ eyes, and discuss their life history strategies. The samples were collected in different seasons in two years (2020 and 2021), with their total length, weight and sex and the number of metacercariae living on them being recorded. The prevalence and mean abundance of infection of different groups and in different seasons were calculated. To find out whether the parasite exhibits preferences for the hosts, a non-parametric test was conducted for independent groups to identify any significant difference in the size of parasite population on male/female hosts, left/right eyes of hosts and different parts of hosts’ eyes. A total of 165 samples [total length: 28.7—49.5 cm; average length: (37.9±4.0) cm; weight: 196.9—827.2 g; average weight: (473.3±127.9) g], including 82 female samples and 83 male samples, were dissected, with 515 parasites being found. The maximum number of parasites observed is 32 worms per fish. In terms of seasonal difference, the prevalence and mean abundance of infection peak in the summer of 2020 and drop in the following autumn and spring. In the summer of 2021, the prevalence of infection is on the decline, while the mean abundance of infection is on the rise. Similar prevalence and mean abundance of infection are observed in the autumn in both 2020 and 2021. To find out group difference, the samples were divided into 5 groups by their total length (TL) at the interval of 5 cm. The minimum values of the prevalence and mean abundance of infection are found in the group of which TL is between 25 cm (inclusive) and 30 cm. Groups of which TL is between 30 cm (inclusive) and 35 cm, between 35 cm (inclusive) and 40 cm, and between 40 cm (inclusive) and 45 cm have similar prevalence and mean abundance of infection. The maximum values of the prevalence and mean abundance of infection are observed in the group of which TL is between 45 cm (inclusive) and 50 cm. The parasite does not exhibit preferences for male/female hosts or left/right eyes of hosts, but they significantly prefers the crystalline lens to the vitreous body. Changes in the size of parasite population are closely related to the seasonal changes, water temperature, bird migration, and snails. The size of parasite population grows with the increase in the total length and eye volume of hosts and frequency of parasite exposure. The reason why the group with TL between 45 cm (inclusive) and 50 cm has higher values of prevalence and mean abundance may lie in the fact that they have a higher risk of parasite exposure because they are larger in size, which enables the hosts to survive from severer infection caused by a larger number of parasites. The preferences exhibited by the parasite in selecting hosts and position of eyes are part of adaptative life history strategies for better transmission and reproduction.
-
-
表 1 匙形双穴吸虫在色林错裸鲤左右眼中的寄生情况
Table 1 Parasitic profile of Diplostomum spathaceum in left or right eye of Gymnocypris selincuoensis
指标Index 左眼Left eye 右眼Right eye 匙形双穴吸虫总数Total number 254 261 感染率Prevalence 49.0 43.0 感染强度Intensity 1—16 1—23 平均丰度Mean abundance 1.5±2.6 1.6±3.2 表 2 匙形双穴吸虫在色林错裸鲤眼球不同部位的寄生情况
Table 2 Parasitic profile of Diplostomum spathaceum in different parts of eyes of Gymnocypris selincuoensis
指标Index 晶状体Len 玻璃体Vitreous humour 匙形双穴吸虫总数Total number 447 68 感染率Prevalence 43.0 8.0 感染强度Intensity 1—17 1—18 平均丰度Mean abundance 1.4±2.5 0.2±1.3 表 3 匙形双穴吸虫在不同性别色林错裸鲤中的寄生情况
Table 3 Parasitic profile of Diplostomum spathaceum in different sex of Gymnocypris selincuoensis
指标Index 雌性Femaln=82 雄性Malen=83 感染率Prevalence 64.0 55.0 感染强度Intensity 1—25 1—31 平均丰度Mean abundance 3.1±5.1 3.1±5.2 -
[1] 中国科学院青藏高原综合科学考察队. 西藏河流与湖泊 [M]. 北京: 科学出版社, 1984: 182-189. Qinghai-Tibet Plateau Comprehensive Scientific Investigation Team, Chinese Academy of Sciences. Rivers and Lakes in Tibet [M]. Beijing: Science Press, 1984: 182-189.
[2] 王苏民, 窦鸿身. 中国湖泊志 [M]. 北京: 科学出版社, 1998: 399. Wang S M, Dou H S. Lakes of China [M]. Beijing: Science Press, 1998: 399.
[3] 武云飞, 吴翠珍. 青藏高原鱼类 [M]. 成都: 四川科学技术出版社, 1992: 506-510. Wu Y F, Wu C Z. The Fishes of the Qinghai-Xizang Plateau [M]. Chengdu: Sichuan Science and Technology Press, 1992: 506-510.
[4] 西藏自治区水产局. 西藏鱼类及其资源 [M]. 北京: 中国农业出版社, 1995: 110-111. Bureau of Water Production in Tibet Autonomous Regio. Fishes and Fish Resources in Xizang, in China [M]. Beijing: China Agriculture Press, 1995: 110-111.
[5] 何德奎, 陈毅峰, 陈自明, 等. 色林错裸鲤性腺发育的组织学研究 [J]. 水产学报, 2001, 25(2): 97-102,188. He D K, Chen Y F, Chen Z M, et al. Histological studies on the gonadal development of Gymnocypris selincuoensis [J]. Journal of Fisheries of China, 2001, 25(2): 97-102,188.
[6] 陈毅峰, 何德奎, 曹文宣, 等. 色林错裸鲤的生长 [J]. 动物学报, 2002, 48(5): 667-676. doi: 10.3321/j.issn:0001-7302.2002.05.013 Chen Y F, He D K, Cao W X, et al. Growth of Slincuo Schizothoracini (Gymnocypris selincuoensis) in Selincuo Lake, Tibeten Plateau [J]. Acta Zoologica Sinica, 2002, 48(5): 667-676. doi: 10.3321/j.issn:0001-7302.2002.05.013
[7] 陈毅峰, 何德奎, 陈宜瑜. 色林错裸鲤的年龄鉴定 [J]. 动物学报, 2002, 48(4): 527-533. doi: 10.3969/j.issn.1674-5507.2002.04.014 Chen Y F, He D K, Chen Y Y. Age discrimination of Selincuo Schizothoracini (Gymnocypris selincuoensis) in Selincuo Lake, Tibeten Plateau [J]. Acta Zoologica Sinica, 2002, 48(4): 527-533. doi: 10.3969/j.issn.1674-5507.2002.04.014
[8] 陈毅峰, 何德奎, 段中华. 色林错裸鲤的年轮特征 [J]. 动物学报, 2002, 48(3): 384-392. doi: 10.3321/j.issn:0001-7302.2002.03.014 Chen Y F, He D K, Duan Z H. Annuli characters of Selincuo Schizothoracine fish (Gymnocypris selincuoensis) in Selincuo Lake, Qinghai-Tibetan Plateau [J]. Acta Zoologica Sinica, 2002, 48(3): 384-392. doi: 10.3321/j.issn:0001-7302.2002.03.014
[9] 尤洋, 周彦锋, 何文辉, 等. 西藏色林错裸鲤肌肉营养品质分析 [J]. 上海海洋大学学报, 2009, 18(2): 2187-2192. You Y, Zhou Y F, He W H, et al. An analysis of the nutritive compositions in the muscle of Gymnocypris selincuoensis [J]. Journal of Shanghai Ocean University, 2009, 18(2): 2187-2192.
[10] 刘军. 色林错裸鲤生活史类型的模糊聚类分析 [J]. 水利渔业, 2006, 26(2): 17-18. doi: 10.3969/j.issn.1003-1278.2006.02.008 Liu J. Fuzzy clustering analysis of life history types of Gymnocypris selincuoensis [J]. Reservoir Fisheries, 2006, 26(2): 17-18. doi: 10.3969/j.issn.1003-1278.2006.02.008
[11] Lee Y I, Seo Mi, Chai J Y. Intestinal flukes recovered from a Herring Gull, Larus argentatus, in the Republic of Korea [J]. The Korean Journal of Parasitology, 2020, 58(1): 81-86. doi: 10.3347/kjp.2020.58.1.81
[12] 朱梦莹, 贾舒安, 王娜, 等. 鱼类寄生复口吸虫研究进展 [J]. 动物医学进展, 2013, 34(5): 103-106. doi: 10.3969/j.issn.1007-5038.2013.05.024 Zhu M Y, Jia S A, Wang N, et al. Progress on Diplostomum in fish [J]. Progress in Veterinary Medicine, 2013, 34(5): 103-106. doi: 10.3969/j.issn.1007-5038.2013.05.024
[13] 孙军. 复口吸虫的研究进展 [J]. 生态科学, 2005, 24(2): 168-172. doi: 10.3969/j.issn.1008-8873.2005.02.018 Sun J. Diplostomum and its progress in advance on ecology [J]. Ecologic Science, 2005, 24(2): 168-172. doi: 10.3969/j.issn.1008-8873.2005.02.018
[14] 王桂堂, 李文祥, 邹红, 等. 淡水鱼类重要寄生虫病诊断手册 [M]. 北京: 科学出版社, 2017: 48-50. Wang G T, Li W X, Zou H, et al. Diagnostic Handbook of Important Parasitic Diseases of Freshwater Fish [M]. Beijing: Science Press, 2017: 48-50.
[15] 李继勋. 鱼病防治关键技术及实用图谱 [M]. 北京: 中国农业大学出版社, 2014: 69-70. Li J X. Key Technologies and Practical Atlas of Fish Disease Control [M]. Beijing: China Agricultural University Press, 2014: 69-70.
[16] 汪开毓, 耿毅, 黄锦炉. 鱼病诊治彩色图谱 [M]. 北京: 中国农业出版社, 2012: 107-108. Wang K Y, Geng Y, Huang J L. Color Atlas of Fish Disease Diagnosis and Treatment [M]. Beijing: China Agriculture Press, 2012: 107-108.
[17] 汪建国. 鱼病学 [M]. 北京: 中国农业出版社, 2013. Wang J G. Ichthyopathology [M]. Beijing: China Agriculture Press, 2013.
[18] Faltýnková A, Sures B, Kostadinova A. Biodiversity of trematodes in their intermediate mollusc and fish hosts in the freshwater ecosystems of Europe [J]. Systematic Parasitology, 2016, 93(3): 283-293. doi: 10.1007/s11230-016-9627-y
[19] 马江霞. 额尔齐斯河鳅科鱼类复口吸虫分类及群落生态学研究 [D]. 乌鲁木齐: 新疆农业大学, 2018. Ma J X. Study on the classification and community ecology of Diplostomidae of Cobitidae in Irtysh River [D]. Urumqi: Xinjiang Agricultural University, 2018.
[20] 朱梦莹. 额尔齐斯河鲈科鱼类复口吸虫研究 [D]. 乌鲁木齐: 新疆农业大学, 2014. Zhu M Y. Research of Diplostomum in the Percidae of the Ergis River [D]. Urumqi: Xinjiang Agricultural University, 2014.
[21] 段成任. 额尔齐斯河凶猛鱼类复口吸虫的流行特点及分类研究 [D]. 乌鲁木齐: 新疆农业大学, 2018. Duan C R. Study on the epidemic characteristics and classification of Diplostomidae of ferocious fish in Irtysh River [D]. Urumqi: Xinjiang Agricultural University, 2018.
[22] 番林古丽·热哈提, 焦丽, 党瑞, 等. 额尔齐斯河白斑狗鱼类复口吸虫的流行病学调查 [J]. 水生态学杂志, 2016, 37(2): 96-100. Parengul, Jiao L, Dang R, et al. Epidemiology of Diplostomum in the eyes of Esox lucius in the Ergis River [J]. Journal of Hydroecology, 2016, 37(2): 96-100.
[23] 唐敏, 廖家红, 覃海芬, 等. 西藏色林错国家级自然保护区鸟类区系和多样性[J]. 绵阳师范学院学报, 2021, 40(8): 57-63. Tang M, Liao J H, Qin H F, et al. On the avifauna and diversity of the Selinko national nature reserve in Tibet [J]. Journal of Mianyang Teachers’ College, 2021, 40(8): 57-63.
[24] 陈毅峰, 陈自明, 何德奎, 等. 藏北色林错流域的水文特征 [J]. 湖泊科学, 2001, 13(1): 21-28. doi: 10.3321/j.issn:1003-5427.2001.01.004 Chen Y F, Chen Z M, He D K, et al. Hydrographic features of Serling Co, north Tibetan Plateau [J]. Journal of Lake Sciences, 2001, 13(1): 21-28. doi: 10.3321/j.issn:1003-5427.2001.01.004
[25] Waadu G D, Chappell L H. Effect of water temperature on the ability of Diplostomum spathaceum miracidia to establish in lymnaeid snails [J]. Journal of Helminthology, 1991, 65(3): 179-185. doi: 10.1017/S0022149X00010671
[26] Lyholt H C K, Buchmann K. Diplostomum spathaceum: effects of temperature and light on cercarial shedding and infection of rainbow trout [J]. Diseases of Aquatic Organisms, 1996, 25(3): 169-173.
[27] Stables J N, Chappell L H. The epidemiology of diplostomiasis in farmed rainbow trout from north-east Scotland [J]. Parasitology, 1986, 92(Pt 3): 699-710.
[28] Stables J N. Studies on the speciation, epidemiology and immunology of Diplostomum spathaceum in fish [D]. Aberdeen: University of Aberdeen, 1984.
[29] Karvonen A, Savolainen M, Seppälä O, et al. Dynamics of Diplostomum spathaceum infection in snail hosts at a fish farm [J]. Parasitology Research, 2006, 99(4): 341-345. doi: 10.1007/s00436-006-0137-8
[30] Hakalahti T, Karvonen A, Valtonen E T. Climate warming and disease risks in temperate regions-Argulus coregoni and Diplostomum spathaceum as case studies [J]. Journal of Helminthology, 2006, 80(2): 93-98. doi: 10.1079/JOH2006351
[31] Karvonen A, Seppälä O, Valtonen E T. Parasite resistance and avoidance behaviour in preventing eye fluke infections in fish [J]. Parasitology, 2004(129): 159-164. doi: 10.1017/S0031182004005505
[32] Sandland G J, Goater C P, Danylchuk A J, et al. Population dynamics of Ornithodiplostomum ptychocheilus metacercariae in fathead minnows (Pimephales promelas) from four northern-Alberta lakes [J]. The Journal of Parasitology, 2001, 87(4): 744-748. doi: 10.1645/0022-3395(2001)087[0744:PDOOPM]2.0.CO;2
[33] Sweeting R A. Investigations into natural and experimental infections of freshwater fish by the common eye-fluke Diplostomum spathaceum Rud [J]. Parasitology, 1974, 69(3): 291-300. doi: 10.1017/S0031182000062995
[34] Burrough R J. The population biology of two species of eyefluke, Diplostomum spathaceum and Tylodelphys clavata, in roach and rudd [J]. Journal of Fish Biology, 1978, 13(1): 19-32. doi: 10.1111/j.1095-8649.1978.tb03409.x
[35] Betterton C. Studies on the host specificity of the eyefluke, Diplostomum spathaceum, in brown and rainbow trout [J]. Parasitology, 1974, 69(1): 11-29. doi: 10.1017/S003118200004614X
[36] Chappell L H. The parasites of the three‐spined stickleback Gasterosteus aculeatus L. from a Yorkshire pond I. seasonal variation of parasite fauna [J]. Journal of Fish Biology, 1969, 1(2): 137-152. doi: 10.1111/j.1095-8649.1969.tb03848.x
[37] Shigin A A. The life-span of Diplostomum spathaceum in the intermediate host [J]. Trudy Gelmintologicheskoi Laboratorii Akademiya Nauk SSSR, 1964(14): 262-272.
[38] 郭爱民, 喀迪尔丁·艾尔肯, 郝翠兰, 等. 新疆虹鳟贝氏复口吸虫的空间分布 [J]. 水生态学杂志, 2020, 41(4): 102-107. doi: 10.15928/j.1674-3075.2020.04.014 Guo A M, Kadierding A, Hao C L, et al. Spatial distribution of Diplostomum baeri in Oncorhynchus mykiss in Xinjiang Province [J]. Journal of Hydroecology, 2020, 41(4): 102-107. doi: 10.15928/j.1674-3075.2020.04.014
[39] 党瑞, 郝翠兰, 焦丽, 等. 额尔齐斯河河鲈复口吸虫的空间分布研究 [J]. 水生态学杂志, 2016, 37(3): 102-106. doi: 10.15928/j.1674-3075.2016.03.015 Dang R, Hao C L, Jiao L, et al. Spatial distribution of Diplostomum in Perca fluviatilis in Ergis River [J]. Journal of Hydroecology, 2016, 37(3): 102-106. doi: 10.15928/j.1674-3075.2016.03.015
[40] Seppälä O, Karvonen A, Valtonen E T. Impaired crypsis of fish infected with a trophically transmitted parasite [J]. Animal Behaviour, 2005, 70(4): 895-900. doi: 10.1016/j.anbehav.2005.01.021
[41] Seppälä O, Karvonen A, Valtonen E T. Manipulation of fish host by eye flukes in relation to cataract formation and parasite infectivity [J]. Animal Behaviour, 2005, 70(4): 889-894. doi: 10.1016/j.anbehav.2005.01.020
[42] Seppälä O, Karvonen A, Valtonen E T. Parasite-induced change in host behaviour and susceptibility to predation in an eye fluke-fish interaction [J]. Animal Behaviour, 2004, 68(2): 257-263. doi: 10.1016/j.anbehav.2003.10.021
[43] Crowden A E, Broom D M. Effects of the eyefluke, Diplostomum spathaceum, on the behaviour of dace (Leuciscus leuciscus) [J]. Animal Behaviour, 1980, 28(1): 287-294. doi: 10.1016/S0003-3472(80)80031-5
[44] Seppälä O, Karvonen A, Valtonen E T. Host manipulation by parasites and risk of non-host predation: is manipulation costly in an eye fluke-fish interaction [J]? Evolutionary Ecology Research, 2006, 8(5): 871-879.
[45] Moore J. Parasites and the Behavior of Animals [M]. New York: Oxford University Press, 2002.
[46] Wayland M T, Chubb J C. A new R package and web application for detecting bilateral asymmetry in parasitic infections [J]. Folia Parasitologica, 2016(63): 2016.039.
[47] Graczyk T. Cases of bilateral asymmetry of Diplostomum pseudospathaceum Niewiadomska, 1984 metacercariae infections (Trematoda: Diplostomidae) in the eye lens of fish [J]. Acta Parasitologica Polonica, 1991(36): 131-134.
[48] Rau M E, Gordon D M, Curtis M A. Bilateral asymmetry of Diplostomum infections in the eyes of lake whitefish Coregonm clupeaformis (Mitchill) and a computer simulation of the observed metacercarial distribution [J]. Journal of Fish Diseases, 1979, 2(4): 291-297. doi: 10.1111/j.1365-2761.1979.tb00171.x
[49] Morley N J, Lewis J W. Influence of Ligula intestinalis plerocercoids (Cestoda: Diphyllobothriidea) on the occurrence of eyeflukes in roach (Rutilus rutilus) from a lake in south-east England [J]. Journal of Helminthology, 2019, 93(1): 66-70. doi: 10.1017/S0022149X17001213
[50] Piete T J J, Koprivnikar J, Orlofske S A, et al. Making the right choice: testing the drivers of asymmetric infections within hosts and their consequences for pathology [J]. Oikos, 2014, 123(7): 875-885. doi: 10.1111/oik.01044