长江上游圆口铜鱼的生物学研究
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摘要
圆口铜鱼(Coreius guichenoti(Sauvage et Dabry))隶属鲤科、鮈亚科、铜鱼属,是长江上游江段的特有鱼类和主要经济鱼类。根据中国科学院水生生物研究所1997年以来的统计,其在上游的渔获物重量组成中通常占据首位。圆口铜鱼是一种河流洄游型鱼类,其整个生活史均在河道中完成。随着栖息地环境变迁和过度捕捞,圆口铜鱼栖息范围萎缩,资源量已经呈显著下降趋势。
本文以长江上游圆口铜鱼为研究对象,对其年龄与生长、繁殖生物学两方面进行了研究,内容涉及圆口铜鱼年龄鉴定、种群结构和生长,以及繁殖生物学等,主要研究方法和结果如下:
2005—2007年,对攀枝花、宜宾、合江、万州、宜昌和宜都几个采样点的渔获物进行分析,共收集到标本7476尾。样本主要是从当地渔民、鱼贩和农贸市场处购得。捕捞网具包括流刺网、小钩和电鱼机等。繁殖生物学研究部分数据和材料来自2004年采集,样本总数1470尾,采集地点包括攀枝花、宜宾、合江、重庆涪陵、重庆木洞、万州和宜昌等站点的长江干流江段。
圆口铜鱼种群年龄结构比较复杂,渔获物由1—7龄共7个龄组组成。优势龄组为1—4龄,占总数的91.88%,5龄及以上数量急剧下降。群体体长分布于22.0—485.0mm之间,体长60—220mm的个体占群体总数的81.97%;体重分布范围为0.1—2100.0g之间,体重0.1—150.0g个体占总数的78.40%。
圆口铜鱼为匀速生长类型。可以用von Bertalanffy方程描述其生长特性:L_t=720.4011[1-e~(-0.1096(t+0.7788))]W_t=4587.6850[1-e~(-0.1096(t+0.7788))]~(3.0311)
生长拐点年龄t=9.34龄,对应体长和体重分别为:L_t=482.73mm,W_t=1363.26g。另估算得出圆口铜鱼极限年龄T_(max)=26龄。
圆口铜鱼初次性成熟年龄为4龄。繁殖季节为4月下旬至7月中旬,以5—6月为盛产期。雌鱼成熟系数在5月达到最大,6月次之,而雄鱼出现5月和7月两个高峰期。对繁殖亲本和早期生活史阶段—卵苗的监测显示,2007年最早发现圆口铜鱼卵粒是在4月22日,采集地点为攀枝花江段,平均水温19.5-20.3℃,平均透明度14.0-15.6cm;而采集到Ⅳ期性腺的最早日期为4月29日,采集地点也在攀枝花江段。
其绝对繁殖力介于4464—131920粒,平均值为46386±51589.28粒,相对繁殖力为5—73粒/g体重,平均值为36.60±27.65粒/g体重。对Ⅳ期卵巢158颗卵母细胞的直径测定的结果显示,卵径范围0.74—2.20mm,平均1.59±0.31mm,峰值出现在1.5—1.7mm,占总卵粒数的33.54%。其卵径分布不均匀,应属于分批产卵类型。
Coreius guichenoti(Sauvage et Dabry),endemic to the upper section of the Yangtze River,belongs to Cypriniformes,Cyprinidae,Rhinogobio.It is one of the main commercial fishes in the upper section of the Yangtze River and weighs most in the catches of the upper section of the Yangtze River.C.guichenoti is a migratory fish,and lives through its whole life history in the river.However,C.guichenoti has a tendency of shrinking in habitat and declining in resource due to the environmental change and overfishing.
This study mainly deals with the following two aspects of C.guichenoti:the age and growth and reproduction.The study also investigates the environment of the major survey stations in the upper section of the Yangtze River.The main results are as follows:
7476 specimens were collected from 2005 to 2007 in Panzhihua,Yibin,Hejiang, Wanzhou,Yichang and Yidu,mainly from fishermen,fishmongers and the markets.The fishing gears include gill nets,crotchets and electro fisher.Partial data obtained during 2004 were adopted and 1470 specimens in total were used for the reproductive biology study,which were collected from Panzhihua,Yibin,Hejiang,Fuling,Mudong,Wanzhou and Yichang.
The C.guichenoti stock had a relatively complex age structure which was composed of seven age groups from one to seven.The preponderant age groups in catch were 1-4 accounting for 91.88%in total and the number of individuals from age 5 upwards declined sharply.The body length in catch ranged from 22.0 to 485.0mm,mainly 60-220mm(81.97%) and the body weight ranged from 0.1-2100.0g,mainly 0.1-150.0g (78.40%).
The growth of C.guichenoti belongs to isometric pattern,which could be described by von Bertalanffy equation.The growth equations of body-length and body-weight were as follows:
L_t=720.4[1-e~(-0.1096(t+0.7788))]W_t=4587.7[1-e~(-0.1096(t+0.7788))]~(3.0311)
The body-weight growth inflexion point is 9.34,with corresponding body-length of 482.73mm,and the body-weight of 1363.26g.The study also calculates the limited age of C.guichenoti(Tmax) which is 26 years old.
C.guichenoti matured at the age of 4 years old and the spawning season was from late April to middle July,with May and June as the reproduction peak period.The GSI of female attained highest during May and then during June.Male have two reproductive peaks:May and July.The spawned eggs of C.guichenoti first occurred on April 22 in 2007 when the average of water temperature was 19.5 to 20.3℃and the average Secchi water transparency was 14.0 to 15.6cm.The gonads at stage IV were first found on April 29 in 2007.All the spawned eggs and mature females were found in Panzhihua.
The absolute fecundity of C.guichenoti is from 4464 to 131920 per fish,with an average of 46386±51589.28.The relative fecundity of C.guichenoti is from 5 to 73 eggs/g body weight,with an average of 36.60±27.65.The diameters of 158 oocytes sampled from a mature female of C.guichenoti were measured It ranges from 0.74 to 2.20mm,with an average of 1.59±0.31mm and 33.54%of oocytes ranged between 1.5 to 1.7mm.It was inferred from the egg-diameter distribution that C.guichenoti might be batch spawners.
本文以长江上游圆口铜鱼为研究对象,对其年龄与生长、繁殖生物学两方面进行了研究,内容涉及圆口铜鱼年龄鉴定、种群结构和生长,以及繁殖生物学等,主要研究方法和结果如下:
2005—2007年,对攀枝花、宜宾、合江、万州、宜昌和宜都几个采样点的渔获物进行分析,共收集到标本7476尾。样本主要是从当地渔民、鱼贩和农贸市场处购得。捕捞网具包括流刺网、小钩和电鱼机等。繁殖生物学研究部分数据和材料来自2004年采集,样本总数1470尾,采集地点包括攀枝花、宜宾、合江、重庆涪陵、重庆木洞、万州和宜昌等站点的长江干流江段。
圆口铜鱼种群年龄结构比较复杂,渔获物由1—7龄共7个龄组组成。优势龄组为1—4龄,占总数的91.88%,5龄及以上数量急剧下降。群体体长分布于22.0—485.0mm之间,体长60—220mm的个体占群体总数的81.97%;体重分布范围为0.1—2100.0g之间,体重0.1—150.0g个体占总数的78.40%。
圆口铜鱼为匀速生长类型。可以用von Bertalanffy方程描述其生长特性:L_t=720.4011[1-e~(-0.1096(t+0.7788))]W_t=4587.6850[1-e~(-0.1096(t+0.7788))]~(3.0311)
生长拐点年龄t=9.34龄,对应体长和体重分别为:L_t=482.73mm,W_t=1363.26g。另估算得出圆口铜鱼极限年龄T_(max)=26龄。
圆口铜鱼初次性成熟年龄为4龄。繁殖季节为4月下旬至7月中旬,以5—6月为盛产期。雌鱼成熟系数在5月达到最大,6月次之,而雄鱼出现5月和7月两个高峰期。对繁殖亲本和早期生活史阶段—卵苗的监测显示,2007年最早发现圆口铜鱼卵粒是在4月22日,采集地点为攀枝花江段,平均水温19.5-20.3℃,平均透明度14.0-15.6cm;而采集到Ⅳ期性腺的最早日期为4月29日,采集地点也在攀枝花江段。
其绝对繁殖力介于4464—131920粒,平均值为46386±51589.28粒,相对繁殖力为5—73粒/g体重,平均值为36.60±27.65粒/g体重。对Ⅳ期卵巢158颗卵母细胞的直径测定的结果显示,卵径范围0.74—2.20mm,平均1.59±0.31mm,峰值出现在1.5—1.7mm,占总卵粒数的33.54%。其卵径分布不均匀,应属于分批产卵类型。
Coreius guichenoti(Sauvage et Dabry),endemic to the upper section of the Yangtze River,belongs to Cypriniformes,Cyprinidae,Rhinogobio.It is one of the main commercial fishes in the upper section of the Yangtze River and weighs most in the catches of the upper section of the Yangtze River.C.guichenoti is a migratory fish,and lives through its whole life history in the river.However,C.guichenoti has a tendency of shrinking in habitat and declining in resource due to the environmental change and overfishing.
This study mainly deals with the following two aspects of C.guichenoti:the age and growth and reproduction.The study also investigates the environment of the major survey stations in the upper section of the Yangtze River.The main results are as follows:
7476 specimens were collected from 2005 to 2007 in Panzhihua,Yibin,Hejiang, Wanzhou,Yichang and Yidu,mainly from fishermen,fishmongers and the markets.The fishing gears include gill nets,crotchets and electro fisher.Partial data obtained during 2004 were adopted and 1470 specimens in total were used for the reproductive biology study,which were collected from Panzhihua,Yibin,Hejiang,Fuling,Mudong,Wanzhou and Yichang.
The C.guichenoti stock had a relatively complex age structure which was composed of seven age groups from one to seven.The preponderant age groups in catch were 1-4 accounting for 91.88%in total and the number of individuals from age 5 upwards declined sharply.The body length in catch ranged from 22.0 to 485.0mm,mainly 60-220mm(81.97%) and the body weight ranged from 0.1-2100.0g,mainly 0.1-150.0g (78.40%).
The growth of C.guichenoti belongs to isometric pattern,which could be described by von Bertalanffy equation.The growth equations of body-length and body-weight were as follows:
L_t=720.4[1-e~(-0.1096(t+0.7788))]W_t=4587.7[1-e~(-0.1096(t+0.7788))]~(3.0311)
The body-weight growth inflexion point is 9.34,with corresponding body-length of 482.73mm,and the body-weight of 1363.26g.The study also calculates the limited age of C.guichenoti(Tmax) which is 26 years old.
C.guichenoti matured at the age of 4 years old and the spawning season was from late April to middle July,with May and June as the reproduction peak period.The GSI of female attained highest during May and then during June.Male have two reproductive peaks:May and July.The spawned eggs of C.guichenoti first occurred on April 22 in 2007 when the average of water temperature was 19.5 to 20.3℃and the average Secchi water transparency was 14.0 to 15.6cm.The gonads at stage IV were first found on April 29 in 2007.All the spawned eggs and mature females were found in Panzhihua.
The absolute fecundity of C.guichenoti is from 4464 to 131920 per fish,with an average of 46386±51589.28.The relative fecundity of C.guichenoti is from 5 to 73 eggs/g body weight,with an average of 36.60±27.65.The diameters of 158 oocytes sampled from a mature female of C.guichenoti were measured It ranges from 0.74 to 2.20mm,with an average of 1.59±0.31mm and 33.54%of oocytes ranged between 1.5 to 1.7mm.It was inferred from the egg-diameter distribution that C.guichenoti might be batch spawners.
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62.Alm G..Connection between maturity,size,and age in fishes.Rep.Inst Freshwat.Res.,1959,40:5-145
63.Casselman J M.Age and growth assessment of fish from their calcified structures-techniques and tools.Noaa Tech.Rep.NMFS,1983,8:1-17
64.Casselman J M.Determination of age and growth.In Weatherley A.H.and Gill,H.S.eds,The biology of Fish growth.Academic Press.1987,209-242
65.Clarke W C,and Blackburn J.Effect of growth on early sexual maturation in stream-type Chinook salmon.Aquaculture,1994,121:95-103
66.Costello M J.Predator feeding strategy and prey importance:A new graphical analysis.J.fish.Bio.,1990,36:261-263
67. Devries D R, and Frie R V. Determination of age and growth. In Murphy B R. and Willis D W., editors. Fisheries techniques (2nd). American Fisheries Society, Bethesda, Maryland. 1996,483-512
68. Fox M G, and Keast A. Effect of overwinter mortality on reproductive life history characteristics of pumpkinseed (Lepomis gibbosus) populations. Can.J.fish. Aquat.Sci, 1991,48:1792-1799
69. Geen G H. Ecological consequences of the proposed Moran Dam on the Fraser River. J. fish. Res. Board Can., 1975, 32:126-135
70. Jones R, and HaII W B. Some observations on the population dynamics of the larval stage in the common gadoids. In Blaxter, J. H. S. editor. The Early life history of Fish. Berlin: Springer-Verlag. 1974: 3-19
71. Klar G T, Parker N C, and Goudie C A. Comparison of growth among families of channel catfish. Prog. Fish.Cult., 1988.50(3): 173-178
72. Kimura D K, Mandapat R R, and Oxford S L. Method, validity, and variability in the age determination of yellowtail rockfish (Sebastes Flavidus), using otoliths. J. Fish. Res. Board. Can., 1979, 36:377-383
73. Laine A O, and Momot W T. Accuracy of using scale and cleithra for aging northern pike from an oligotrophic Onatario Lake. North Am. J. fish. Manag., 1991, 11:220-225
74. Mann R H K, and Mills C A. Variations in the size of gonads, eggs and larvae of the dace. Leuciscus Leuciscus. Env. Biol. Fish., 1985, 13:277-287
75. Mann R H K, Mills C A, and Crisp D T. Geographic variation in the life history tactics of some species of freshwater fish. In Potts G W. and Wotton R J. editors. Fish reproduction: strategies and tactics. Academic Press, London, England, 1984: 171-186
76. Michael R O', and Botsford L W. Estimating the status of neat shore rockfish (Sebastes spp.) populations with length frequency data. Eco. App., 2006, 16(3):977-986
77. Pannella G. Fish otoliths; daily growth layers and periodical patterns. Science 1971, 173: 1124-1127
78. Renick D A, Bryga H, and Endler J A. Experimentally induced life-history evolution in a natural population. Nature, 1990, 346:357-359
79. Ricker W E. Changes in the average size and average age of Pacific salmon. Can. J. Fish. Aquat.Sci, 1981, 38:1636-1656
80. Rowe D K, and Thorpe J E. Suppression of maturation of maturation in male Atlantic Salmon {Salmon salar L.) Parr by reduction in feeding and growth during spring months. Aquaculture, 1990, 86:291-313
81. Saunders R L, and Henderson E B. Effects of constant daylength on sexual maturation and growth of Atlantic salmon {Salmon salar L.) parr. Can. J. Fish. Aquat. Sci, 1988,45:60-64
82. Silverstein J T, and Shimma H. Effect of restricted feeding on early maturation in female and male amago salmon, Oncorhynchus masou ishikawae. J. Fish Biol, 1994, 45:1133-1135
83. Stearns S C. A natural experiment in life-history evolution; field data on the introduction of mosquito fish {Grabuia affinis) to Hawii. Evoltion, 1983, 37:266-281
84. Stock assessment for tropical small-scale fisheries. Proceedings of an international workshop held at the University of Rhode Island. Int. Cent. Mar. Res. Dev., University of Rhode Island. Kingston. 1979, 9: 19-21
85. Thompson K P. and Beckman D W. Validation of age estimates from white sucker otoliths. Trans. Am. Fish. Aquat. Sci., 1995,44:1485-1495
86. Ware D M. Relation between egg size, growth, and natural mortality of larva fish. J. Fish. Res. Bd Can, 1975, 32:2503-2513
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55.中国科学院水生生物研究所.长江三峡工程生态与环境监测系统鱼类及水生生物重点站2007年度报告.2008
56.中国科学院水生生物研究所.金沙江下游和长江上游干流鱼类资源调查与现状评价报告.1998
57.郑曙明,吴青.铜鱼和圆口铜鱼耗氧率的研究.四川畜牧兽医学院学报,1998,12(3-4):6-8
58.郑曙明.铜鱼和圆口铜鱼肌肉氨基酸的初步研究.四川畜牧兽医学院学报,1998,12(3-4):23-25
59.郑曙明,熊全沫.铜鱼卵巢的显微和超微结构的研究.武汉大学学报:自然科学版,1993,39(3):103-109
60.庄平,曹文宣.长江中、上游铜鱼的生长特性.水生生物学报,1999,23(6):577-583
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63.Casselman J M.Age and growth assessment of fish from their calcified structures-techniques and tools.Noaa Tech.Rep.NMFS,1983,8:1-17
64.Casselman J M.Determination of age and growth.In Weatherley A.H.and Gill,H.S.eds,The biology of Fish growth.Academic Press.1987,209-242
65.Clarke W C,and Blackburn J.Effect of growth on early sexual maturation in stream-type Chinook salmon.Aquaculture,1994,121:95-103
66.Costello M J.Predator feeding strategy and prey importance:A new graphical analysis.J.fish.Bio.,1990,36:261-263
67. Devries D R, and Frie R V. Determination of age and growth. In Murphy B R. and Willis D W., editors. Fisheries techniques (2nd). American Fisheries Society, Bethesda, Maryland. 1996,483-512
68. Fox M G, and Keast A. Effect of overwinter mortality on reproductive life history characteristics of pumpkinseed (Lepomis gibbosus) populations. Can.J.fish. Aquat.Sci, 1991,48:1792-1799
69. Geen G H. Ecological consequences of the proposed Moran Dam on the Fraser River. J. fish. Res. Board Can., 1975, 32:126-135
70. Jones R, and HaII W B. Some observations on the population dynamics of the larval stage in the common gadoids. In Blaxter, J. H. S. editor. The Early life history of Fish. Berlin: Springer-Verlag. 1974: 3-19
71. Klar G T, Parker N C, and Goudie C A. Comparison of growth among families of channel catfish. Prog. Fish.Cult., 1988.50(3): 173-178
72. Kimura D K, Mandapat R R, and Oxford S L. Method, validity, and variability in the age determination of yellowtail rockfish (Sebastes Flavidus), using otoliths. J. Fish. Res. Board. Can., 1979, 36:377-383
73. Laine A O, and Momot W T. Accuracy of using scale and cleithra for aging northern pike from an oligotrophic Onatario Lake. North Am. J. fish. Manag., 1991, 11:220-225
74. Mann R H K, and Mills C A. Variations in the size of gonads, eggs and larvae of the dace. Leuciscus Leuciscus. Env. Biol. Fish., 1985, 13:277-287
75. Mann R H K, Mills C A, and Crisp D T. Geographic variation in the life history tactics of some species of freshwater fish. In Potts G W. and Wotton R J. editors. Fish reproduction: strategies and tactics. Academic Press, London, England, 1984: 171-186
76. Michael R O', and Botsford L W. Estimating the status of neat shore rockfish (Sebastes spp.) populations with length frequency data. Eco. App., 2006, 16(3):977-986
77. Pannella G. Fish otoliths; daily growth layers and periodical patterns. Science 1971, 173: 1124-1127
78. Renick D A, Bryga H, and Endler J A. Experimentally induced life-history evolution in a natural population. Nature, 1990, 346:357-359
79. Ricker W E. Changes in the average size and average age of Pacific salmon. Can. J. Fish. Aquat.Sci, 1981, 38:1636-1656
80. Rowe D K, and Thorpe J E. Suppression of maturation of maturation in male Atlantic Salmon {Salmon salar L.) Parr by reduction in feeding and growth during spring months. Aquaculture, 1990, 86:291-313
81. Saunders R L, and Henderson E B. Effects of constant daylength on sexual maturation and growth of Atlantic salmon {Salmon salar L.) parr. Can. J. Fish. Aquat. Sci, 1988,45:60-64
82. Silverstein J T, and Shimma H. Effect of restricted feeding on early maturation in female and male amago salmon, Oncorhynchus masou ishikawae. J. Fish Biol, 1994, 45:1133-1135
83. Stearns S C. A natural experiment in life-history evolution; field data on the introduction of mosquito fish {Grabuia affinis) to Hawii. Evoltion, 1983, 37:266-281
84. Stock assessment for tropical small-scale fisheries. Proceedings of an international workshop held at the University of Rhode Island. Int. Cent. Mar. Res. Dev., University of Rhode Island. Kingston. 1979, 9: 19-21
85. Thompson K P. and Beckman D W. Validation of age estimates from white sucker otoliths. Trans. Am. Fish. Aquat. Sci., 1995,44:1485-1495
86. Ware D M. Relation between egg size, growth, and natural mortality of larva fish. J. Fish. Res. Bd Can, 1975, 32:2503-2513
87. Wotton R J. Ecology of Teleost Fishes. New York: Chapman and Hall Ltd., 1990.