THEORIES OF FISH ATTRACTION AND REPULSION IN RIVERS AND THEIR APPLICATIONS IN FISH PASSAGE SYSTEMS
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摘要:
过鱼设施作为恢复鱼类洄游通道的重要生态补偿措施, 已广泛应用于水生态保护, 然而其过鱼效果尚未达到预期。诱驱鱼技术是提升过鱼设施过鱼效果的关键手段之一。结合近十年的室内与野外实验研究, 文章系统阐述了水流、声、光、电、气泡幕及生物因子等对鱼类行为的影响机制及诱驱理论, 总结了诱驱鱼技术在过鱼设施中的应用优势与不足, 得出结论: (1)鱼类对诱驱鱼因素的行为响应研究逐渐深入, 由二维平面发展到三维空间(如由流速到流场、由声强到声场等); (2)诱驱鱼因素对不同鱼类的诱驱效果呈现种属共性和特异性, 且不同大小鱼类对诱驱鱼因素也存在行为响应差异; (3)诱驱鱼技术研究对象逐渐由海洋鱼类转向淡水经济鱼类和保护鱼种, 研究方法趋向于考虑多因子交互作用对鱼类行为的影响; (4)诱驱鱼技术的效果受多种因素交互影响, 包括物种差异、工程布置适应性、运行管理等。未来研究应进一步解析多因子交互作用下的鱼类行为响应机制, 优化适应不同过鱼设施运行条件的诱驱鱼设计, 推动水生态保护的发展。
Abstract:Fish passage facilities, as an important ecological compensation measure for restoring fish migratory routes, have been widely implemented in aquatic ecosystem conservation. However, their effectiveness in facilitating fish passage remains below expectations. Fish attraction and repulsion technologies play a critical role in enhancing fish passage performance. This paper systematically elucidates the mechanisms and guidance theories of water flow, sound, light, electricity, bubble curtains, and biological factors on fish behavior, based on nearly a decade of indoor and field experiments. The paper also summarizes the advantages and limitations of these technologies in fish passage applications. Key conclusions include: (1) Research on fish behavioral responses to attraction and repulsion factors has evolved from two-dimensional to three-dimensional perspectives (e.g., from flow velocity to flow fields, from sound intensity to sound fields); (2) The effectiveness of attraction and repulsion factors shows both commonalities and species-specific differences among fish, with varying behavioral responses across different fish sizes; (3) Research focus has shifted from marine species to freshwater economic and protected species, with increasing attention to the interactive effects of multiple factors on fish behavior; And (4) the performance of attraction and repulsion technologies is influenced by multiple interacting factors, including species-specific characteristics, engineering design adaptability, and operational management. Future research should further explore fish behavioral responses under multi-factor interactions, optimize attraction and repulsion designs for various operational conditions of fish passage facilities, and advance aquatic ecosystem conservation.
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拒马河原名涞水河, 是海河流域大清河北部支流, 发源于河北省涞源县西北太行山麓, 流经紫荆关向北至涞水县折向东流, 至北京市张坊镇分为南、北二支, 南拒马河目前已经干涸消亡, 北拒马河于白沟镇流入大清河, 拒马河干流长254 km, 张坊镇以上流域面积5115 km2, 是中国北方最大的冬季不结冰河流[1—4]。拒马河流域地貌复杂, 不同河段气候与环境相差悬殊, 生境的多样性造就了丰富的水生生物资源, 是华北地区内陆水域生物多样性的重要组成部分, 具有极高的科研及保护价值[5, 6]。
河流及湖泊水文环境的改变影响其中鱼类群落的结构组成, 而鱼类群落的结构组成也是水生生态系统健康的重要指标, 可通过鱼类群落的变动来分析水环境状况的受污染程度[7, 8]。近年来, 拒马河生态环境发生了巨大变化, 生态流量逐年下降, 水生态环境受人类影响较大[5], 水质、水文流通性发生改变, 改变了生活在其中的鱼类的栖息环境, 增加了鱼类的生存压力[9]。拒马河拦河坝的建设导致河流片段化严重, 纵向连通几乎丧失, 鱼类洄游通道被阻断, 部分河段形成小型静水湖泊, 洄游性、亲流性和因拦河坝建设而失去栖息及越冬场所的鱼类减少而喜静水性鱼类增加[10, 11]。加深鱼类群落在空间和时间上对生境改变做出反应的认知对制定有效的生态环境保护策略非常重要[12], 国内关于鱼类群落与环境因子的关系的研究多集中于近海和河口, 对于内陆河流的研究较为匮乏[13—15]。目前, 拒马河鱼类资源的调查虽有不少资料, 但了解得仍然不够深入和系统, 多仅限于北京段, 如2008年杨文波等[16]对拒马河北京段的鱼类资源调查, 记录鱼类24种; 张春光等[17]在《北京及其邻近地区的鱼类》中整理历史资料, 记录拒马河历史上出现鱼类42种; 袁立来等[6]利用鱼类生物完整性指数对拒马河北京段进行了河流健康评价, 发现拒马河北京段河流健康整体处于较差水平。对拒马河上游区域的物种组成和地理分布等基本问题仍缺乏准确的数据, 因此, 系统开展拒马河鱼类资源调查, 以掌握拒马河鱼类时空分布及影响因子, 是拒马河鱼类资源开发利用和资源保护首要解决的问题。
本研究基于2019—2021年间6次拒马河鱼类资源调查数据, 从物种组成、优势种及群落结构特征等方面分析了鱼类群落结构现状, 并讨论了鱼类资源的时空分布与环境因子的关系, 为拒马河流域内的鱼类资源的保护、修复和管理提供理论基础和科学依据。
1. 材料与方法
1.1 样品采集
本次调查范围自涞源县拒马河源头至张坊镇龙安大桥, 采样点的选择参考《内陆水域渔业自然资源调查规范》[18], 综合考虑拒马河河流特点及采样点的代表性, 设置了15个采样点, 采样点的经纬度和位置见表 1和图 1。于2019年5月 (春季)、8月 (夏季)、10月 (秋季), 2020年8月、10月和2021年5月进行采样调查, 每个采样点设置两条多目刺网和地笼, 刺网规格20 m×1 m和25 m×1.5 m, 网目为3和5 cm, 地笼规格为10 m×35 cm×30 cm和10 m×25 cm×20 cm, 网目为1 cm。刺网和地笼于16:00—18:00时下网, 次日5:00—7:00起网, 放置时间约12h。依据最新鱼类学专著对采集的渔获物分类[19—21], 对新鲜鱼类样本现场进行物种鉴定和个体测量, 采用游标卡尺和电子天平测量体长(精确到1 mm)和体重(精确到0.1 g)数据, 现场无法鉴定的个体用95%的酒精保存后带回实验室进一步分类鉴定。
表 1 拒马河采样点分布Table 1. Distribution of sampling stations in the Juma River采样点
Sampling station编号
Serial number经度
Longitude纬度
Latitude拒马河源头
The source of Juma RiverS1 114°45′04.115″ 39°19′57.115″ 刁江汇
DiaojianghuiS2 115°01′36.775″ 39°25′39.702″ 紫荆关大桥
Zijingguan BridgeS3 115°10′03.338″ 39°25′41.776″ 清凉涧
QingliangjianS4 115°15′38.952″ 39°37′44.969″ 小丰口桥
Xiaofengkou BridgeS5 115°20′35.686″ 39°42′05.548″ 别岸
Bie’anS6 115°27′38.115″ 39°39′06.187″ 琅琊河
Langya RiverS7 115°29′18.773″ 39°39′08.849″ 天花板
TianhuabanS8 115°30′40.557″ 39°39′47.696″ 北石门
BeishimenS9 115°32′44.611″ 39°38′33.555″ 西河口
XihekouS10 115°34′09.356″ 39°38′27.367″ 九渡Jiudu S11 115°35′22.049″ 39°37′41.080″ 六渡Liudu S12 115°37′59.485″ 39°38′19.132″ 穆家口
MujiakouS13 115°39′48.970″ 39°37′43.089″ 千河口
QianhekouS14 115°39′57.071″ 39°36′38.520″ 龙安大桥
Longan BridgeS15 115°41′14.921″ 39°34′32.855″ 对鱼类资源调查的同时, 对15个采样点进行海拔(ASL)、水温(Tem)、水深(Dep)、溶解氧(DO)、透明度(SD)、浊度(FNU)、pH和叶绿素a(Chl.a)等环境因子的测定(水科院, 未发表数据)。
1.2 数据处理分析
采用以下多样性指数来分析拒马河鱼类群落多样性: (1)Margalef丰富度指数(DMa)[22]: DMa =(S–1)/ln N; (2)Shannon-Wiener多样性指数(H') [23]: H′= –∑Pi× ln Pi; (3)Pielou均匀度指数(J′) [24]: J′=H′/lnS。式中, S为采集到的鱼类种类数, N为采集到鱼类的个体数, Pi为样品中第i种鱼类的个体数占渔获物中全部个体数的比例。选择Pinkas等[25]提出的相对重要指数(Index of relative importance, IRI)计算拒马河鱼类群落优势种: IRI=(Ni+Wi) Fi。式中, Ni为渔获物中第i种鱼类的个体数占渔获物总个体数的百分比, Wi为第i种鱼类的生物量占渔获物总生物量的百分比, Fi为第i种鱼类在采样点出现的频率。定义IRI≥500的为优势种, 100≤IRI<500的为重要种, 10≤IRI<100的为常见种, 1≤IRI<10的为一般种, IRI<1的为稀有种[26]。
采用丰度生物量比较曲线(Abundance-biomass comparison curves, ABC曲线) [27]判断鱼类群落稳定性及受干扰程度, ABC曲线的统计量用W表示, 当生物量优势度曲线位于丰度优势度曲线之上时, W为正, 表明群落结构稳定未受干扰, 反之W为负[28]: W=∑(Bi–Ai)/50(S–1)。式中, Ai为第i种鱼类相对应的丰度累积百分比, Bi为第i种鱼类相对应的生物量累积百分比, S为物种数。
为满足数据的正态齐性和方差齐性, 分析前对鱼类丰度数据及多样性数据进行lg(x+1)对数转换, 运用SPSS 25.0通过单因素方差分析(One-way ANOVE)检验不同季节、海拔鱼类丰度及多样性的差异。若存在显著性差异, 进一步使用SNK(Student-newman-Keuls)多重比较分析不同季节、海拔间鱼类丰度及多样性的变化[33]。
在Primer 5.0软件中, 以采集到的鱼类丰度数据为原始矩阵, 进行lg(x+1)对数转换, 运用等级聚类分析(Cluster)和非参数多变量排序(NMDS)将鱼类群落划分不同的组分。运用相似性百分比分析(Similarity percentages, SIMPER)确定维持不同组分间结构相似性的关键物种[29]。
利用Canoco 5.0软件对15个采样点的鱼类物种组成和环境因子进行去趋势对应分析(Detrended correspondence analysis, DCA), 根据分析结果中各排序轴的大小选择线性模型(Redundancy analysis, RDA)或单峰模型(Canonical correspondence analysis, CCA)分析鱼类物种和环境因子的相关性[30]。如果排序轴大于4选择CCA分析, 排序轴小于3选择RDA分析, 介于3和4之间, 两种分析方法均可。
2. 结果
2.1 拒马河鱼类组成及优势种
2019—2021年在拒马河共采集鱼类5486尾, 隶属5目11科37种 (表 2)。渔获物中鲤形目占绝对优势, 有3科24种, 占总种数的64.86%; 鲈形目3科6种, 占16.22%; 鲇形目2科4种, 占10.81%; 合鳃鱼目2科2种, 占5.41%; 颌针鱼目1科1种, 占2.70%。
表 2 拒马河鱼类组成、各月份相对重要指数(IRI)及生态类型Table 2. The fish composition, index of relative important (IRI) of each month and ecological types in the Juma River物种Species 相对重要指数IRI 生态类型 2019.5 2019.8 2019.10 2020.8 2020.10 2021.5 Ecological types 鲤形目Cypriniformes 鲤科Cyprinidae 鲤亚科Cyprininae 鲫Carassius auratus 209.63 3024.43 371.53 1050.27 245.62 273.74 S, Omn, De, V 鱊亚科Acheilognathinae 兴凯鱊Acheilognathus chankaensis 8.36 — — — — — S, Omn, U,Sp 中华鳑鲏Rhodeus sinensis 8.08 — — 3.87 — — S, Omn, U, Sp 高体鳑鲏Rhodeus ocellatus 39.02 148.43 — 62.34 — 4.60 S, Omn, U, Sp 雅罗鱼亚科Leuciscinae 尖头鱥Rhynchocypris oxycephalus 668.20 216.15 101.76 476.61 44.59 90.97 R, Omn, Lo, Dr 拉氏鱥Rhynchocypris lagowskii 317.45 10.61 450.36 54.01 215.97 108.09 R, Omn, Lo, Dr 襁亚科Danioninae 宽鳍鱲Zacco platypus 1928.30 494.22 282.76 2497.27 1558.08 413.94 R, Omn, U, D 马口鱼Opsariichthys bidens 5.48 4.53 14.86 235.48 — — R, Car, U, D 亚科Gobioninae 麦穗鱼Pseudorasbora parva 2291.93 3463.52 1546.54 2742.93 2605.65 5382.96 S, Omn, De, V 点纹银Squalidus wolterstorffi 1162.26 121.92 24.68 92.90 1.49 22.05 S, Omn, Lo, Dr 中间银Squalidus intermedius — 53.15 — 34.37 — — R, Omn, Lo, Dr 兴隆山小鳔Microphysogobio hsinglungshanensis 217.79 53.69 29.31 — 12.42 369.75 R, Omn, Lo, Dr 黑鳍鳈Sarcochilichthys nigripinnis 548.01 622.04 198.44 685.56 383.17 884.47 S, Omn, Lo, Sp 棒花Gobio rivuloides 1.47 183.58 — — — 7.15 R, Omn, De, V 棒花鱼Abbottina rivularis 263.72 38.68 385.70 275.44 532.21 — R, Omn, De, V 蛇Saurogobio dabryi 0.82 1.83 26.13 40.43 337.43 46.09 R, Omn, Lo, Dr 花鳅科Cobitidae 泥鳅Misgurnus anguillicaudatus 1044.35 580.91 55.37 41.34 16.14 50.20 S, Omn, De, Dr 北方泥鳅Misgurnus bipartitus — 1.77 — — — — R, Omn, De, Dr 大鳞副泥鳅Paramisgurnus dabryanus 109.14 144.85 19.20 53.78 21.18 12.62 S, Omn, De, Dr 花斑花鳅Cobitis melanoleuca 95.38 115.57 — 12.07 1.72 71.63 S, Omn, De, Dr 条鳅科Nemacheilidae 北鳅Lefua costata — — — — 2.28 1.41 R, Omn, De, Dr 赛丽高原鳅Triplophysa sellaefer 11.33 — 5.83 — 104.93 37.93 R, Omn, De, Dr 尖头高原鳅TriplophysaCuneicephala — — 2.16 — — — R, Omn, De, Dr 达里湖高原鳅Triplophysa dalaica 97.01 12.11 80.15 — 168.26 127.34 R, Omn, De, Dr 合鳃鱼目Symbranchiformes 刺鳅科Mastacembelidae 刺鳅Sinobdella sinensis 50.12 26.09 — 59.92 — 2.85 S, Car, De, D 合鳃鱼科Symbranchidae 黄鳝Monopterus albus 12.24 — — — — — S, Car, De, Dr 鲈形目Percoidei 鰕虎鱼科Gobiidae 子陵吻鰕虎鱼Rhinogobius giurinus 2.65 8.25 4.56 53.91 38.20 5.10 R, Car, De, D 林氏吻鰕虎鱼Rhinogobius lindbergi 68.08 19.58 9.97 — 0.01 26.13 R, Car, De, D 波氏吻鰕虎鱼Rhinogobius cliffordpopei 44.63 0.96 1.70 — 1.31 — R, Car, De, D 福岛吻鰕虎鱼Rhinogobius fukushimai 30.47 0.63 0.43 1.83 — 2.40 R, Car, De, D 沙塘鳢科Odontobuidae 小黄䱂鱼Micropercops swinhonis 1250.14 308.40 786.30 453.05 386.69 663.59 S, Omn, De, D 丝足鲈科Osphronemidae 圆尾斗鱼Macropodus chinensis 1.40 — 0.44 0.98 1.42 2.62 S, Car, Lo, Dr 鲇形目Siluriformes 鲿科Bagridae 黄颡鱼Pelteobagrus fulvidraco 313.28 861.57 449.50 1265.35 191.14 10.27 S, Car, De, D 瓦氏黄颡鱼Pelteobagrus vachellii 7.92 37.05 0.58 — 9.54 — R, Car, De, D 乌苏里黄颡鱼Pelteobagrus ussuriensis — 287.03 — 37.59 — 20.41 R, Car, De, D 鲇科Siluridae 鲇Silurus asotus 10.83 203.88 198.89 26.22 — — S, Car, De, V 颌针鱼目Beloniformes 青鳉科Adrianichthyidae 青鳉Oryzias sinensis — 1.13 — — — — S, Omn, U, V 注: S. 喜缓流或静水; R. 亲流性; Car. 肉食性; Omn. 杂食性; U. 中上层; Lo. 中下层; De. 底栖。V. 黏性卵; D. 沉性卵; Dr. 漂流性卵; Sp. 喜贝类性卵Note: S. slow flow; R. riffle; Car. carnivore; Omn. omnivore; U. upper; Lo. lower; De. demersal; V. viscid egg; D. demersal egg; Dr. drifting egg; Sp. spawning in shellfish 鱼类生态类型(表 2): 按生活习性可以将采集到的鱼类划分为亲流性鱼类(R)和喜缓流或静水性鱼类(S), 其中亲流性鱼类有20种, 占采集到的鱼类总种数的54.05%, 喜缓流或静水性鱼类有17种, 占总种数的45.95%; 按食性可以划分为杂食性鱼类(Omn)和肉食性鱼类(Car), 杂食性鱼类有25种, 占总种数的67.57%, 肉食性鱼类有12种, 占总种数的32.43%; 按不同栖息水层划分, 底栖鱼类(De)有23种, 占62.16%, 中下层鱼类(Lo)有8种, 占21.62%, 中上层鱼类(Up)有6种, 占16.22%; 按产卵类型划分, 产漂流性卵的鱼类(Dr)有16种, 占43.24%, 产沉性卵的鱼类(D)有11种, 占29.73%, 产黏性卵的鱼类(V)6种, 占16.22%, 喜贝类性产卵的鱼类(Sp)4种, 占10.81%。
以相对重要指数IRI≥500的为优势种, 在6次采样调查中, 拒马河优势种有10种, 重要种有12种, 主要种有8种, 一般种有7种。各月份优势种存在差异, 2019年5月优势种为尖头鱥Rhynchocypris oxycephalus Sauvage、宽鳍鱲Zacco platypus Temminck & Schlegel、麦穗鱼Pseudorasbora parva Temminck & Schlegel、点纹银Squalidus wolterstorffi Regan、黑鳍鳈Sarcochilichthys nigripinnis Günther、泥鳅Misgurnus anguillicaudatus Cantor、小黄䱂鱼Micropercops swinhonis Günther; 2019年8月优势种为鲫Carassius auratus Linnaeus、麦穗鱼、黑鳍鳈、泥鳅、黄颡鱼Tachysurus fulvidraco Richardson; 2019年10月优势种为麦穗鱼、小黄䱂鱼; 2020年8月优势种为鲫、宽鳍鱲、麦穗鱼、黑鳍鳈、黄颡鱼; 2020年10月优势种为宽鳍鱲、麦穗鱼、棒花鱼Abbottina rivularis Basilewsky; 2021年5月优势种为麦穗鱼、黑鳍鳈、小黄䱂鱼。麦穗鱼在6次采样调查中均作为优势种出现。
2.2 拒马河鱼类群落特征
单因素方差分析显示(表 3), 不同季节间个体数(N)、物种数(S)、Shannon-Wiener多样性指数(H')、Margalef丰富度指数(DMa)呈现显著性差异(P<0.05), Pielou均匀度指数(J')无显著性差异(P>0.05)。SNK多重比较分析显示, 物种数、个体数均以2019年5月最多, 15个采样点平均捕获鱼类91.00尾9.67种, 2021年10月最少, 平均38.73尾5.07种; Shannon-Wiener多样性指数(H')、Margalef丰富度指数(DMa)均以2019年8月份最高, 分别是1.76(1.09—2.20)和2.06(1.12—2.98)。2019—2021年调查显示拒马河夏季的Shannon-Wiener多样性指数(H')、Margalef丰富度指数(DMa)和Pielou均匀度指数(J')均值高于春、秋两季(图 2)。
表 3 基于单因素方差分析检验(F值)拒马河鱼类多样性指数不同季节、不同海拔的变化Table 3. Changes of fish diversity index in different seasons and different altitudes in the Juma River based on One-way ANOVA test (F value)项目Item 个体数
Number of
individuals (N)物种数
Number of
species (S)多样性指数
Shannon-Wiener
diversity (H')均匀度指数
Pielou
evenness (J')丰富度指数
Margalef
richness (Dma)季节Season 2.935* 7.986** 6.504** 1.022ns 4.9** 海拔Altitude 12.206** 0.122ns 1.616ns 7.368ns 3.765ns 注: ns、*和**分别代表P>0.05、P<0.05、和P<0.01Note: ns, * and ** represent P>0.05, P<0.05 and P<0.01 ![]()
图 2 拒马河鱼类多样性指数在不同季节、不同海拔差异Figure 2. Differences of fish diversity index in different seasons and different altitudes in the Juma River物种数(S)、Shannon-Wiener多样性指数(H')、Margalef丰富度指数(DMa)、Pielou均匀度指数(J')在不同海拔无显著性差异(P>0.05), 只有个体数(N)呈现显著性差异(P<0.05), 海拔高于500 m的位点平均可采集鱼类143.91尾, 海拔低于500 m的位点可采集48.19尾(图 2)。
NMDS排序分析和聚类分析结果一致, 拒马河鱼类群落自上游至下游可划分为3组, 即S1和S2为组Ⅰ, S3和S4为组Ⅱ, 其余采样点为组Ⅲ(图 3)。相似性百分比分析SIMPER(表 4)显示, 组Ⅰ平均相似度为51.01%, 主要贡献种有麦穗鱼、鲫、小黄䱂鱼、泥鳅、拉氏鱥Rhynchocypris lagowskii Dybowski、赛丽高原鳅Triplophysa sellaefer Nichols、尖头鱥、达里湖高原鳅Triplophysa dalaica Kessler和林氏吻鰕虎鱼Rhinogobius lindbergi Berg, 累积贡献度为90.07%; 组Ⅱ平均相似度为66.92%, 主要贡献种有麦穗鱼、小黄䱂鱼、棒花鱼、黄颡鱼、鲫、子陵吻鰕虎鱼Rhinogobius giurinus Rutter、中间银Squalidus intermedius Nichols和林氏吻鰕虎鱼, 累积贡献度为90.16%; 组Ⅲ平均相似度为61.00%, 主要贡献种有麦穗鱼、宽鳍鱲、小黄䱂鱼、黑鳍鳈、鲫、黄颡鱼、泥鳅、棒花鱼、高体鳑鲏Rhodeus ocellatus Kner、兴隆山小鳔Microphysogobio hsinglungshanensis Mori、点纹银、刺鳅Sinobdella sinensis Bleeker和棒花Gobio rivuloides Nichols, 累积贡献度为91.23%。
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图 3 拒马河鱼类群落Cluster聚类和NMDS排序图Figure 3. Clustering and NMDS sequencing analysis of fish community structure in the Juma River表 4 拒马河不同位点的主要贡献种相似性百分比分析Table 4. The main contributionspecies of Juma River in different locals of similarity percentage analysis物种Species 组Ⅰ GroupⅠ 组Ⅱ GroupⅡ 组Ⅲ GroupⅢ AA AS (%) Con (%) AA AS (%) Con (%) AA AS (%) Con (%) 麦穗鱼Pseudorasbora parva 277.50 10.87 21.30 145.00 14.29 21.36 51.91 9.14 14.99 鲫Carassius auratus 73.00 8.74 17.14 9.00 6.23 9.31 13.09 5.24 8.59 小黄䱂鱼Micropercops swinhonis 24.50 6.56 12.86 41.00 10.97 16.40 31.91 7.28 11.93 泥鳅Misgurnus anguillicaudatus 23.50 6.00 11.76 9.09 3.63 5.95 拉氏鱥Rhynchocypris lagowskii 78.00 3.65 7.16 赛丽高原鳅Triplophysa sellaefer 17.50 2.82 5.54 尖头鱥Rhynchocypris oxycephalus 135.00 2.82 5.54 达里湖高原鳅Triplophysa dalaica 78.50 2.24 4.39 林氏吻鰕虎鱼Rhinogobius lindbergi 3.50 2.24 4.39 3.00 3.29 4.92 棒花鱼Abbottina rivularis 39.50 9.39 14.04 4.73 2.43 3.98 黄颡鱼Pelteobagrus fulvidraco 15.00 7.18 10.73 6.91 3.74 6.12 子陵吻鰕虎鱼Rhinogobius giurinus 8.50 4.82 7.20 中间银Squalidus intermedius 9.00 4.15 6.21 宽鳍鱲Zacco platypus 55.91 8.07 13.22 黑鳍鳈Sarcochilichthys nigripinnis 32.55 7.26 4.33 高体鳑鲏Rhodeus ocellatus 5.55 2.31 3.79 兴隆山小鳔Microphysogobio hsinglungshanensis 13.82 2.16 3.54 点纹银Squalidus wolterstorffi 18.27 2.11 3.45 刺鳅Sinobdella sinensis 2.45 1.21 1.99 棒花Gobio rivuloides 2.36 1.09 1.78 注: AA. 平均多度, AS. 平均相似度, Con. 贡献度Note: AA. average abund, AS. average similarity, Con. contribution 2.3 丰度生物量比较曲线
鱼类群落的优势度曲线变化趋势(图 4)显示, 2020年8月拒马河鱼类群落的生物量优势度曲线位于丰度优势度曲线之上, 统计值W＞0, 表示该月份鱼类群落结构稳定未受干扰, 其余月份的丰度累积百分比与生物量累积百分比的大小交错分布, ABC曲线中丰度优势度曲线与生物量优势度曲线相交, 统计量W＜0, 鱼类群落受到中等程度干扰。2020年夏季数量优势度曲线和丰度优势度曲线均排在前5位的优势种类为麦穗鱼、宽鳍鱲和尖头鱥, 该季度所采集渔获物中, 平均个体生物量为8.99 g, 平均个体生物量最大的是黄颡鱼, 为30.86 g, 平均个体生物量最小的是福岛吻鰕虎鱼Rhinogobius fukushimai Mori, 为0.50 g。
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图 4 拒马河不同季节鱼类群落ABC曲线Figure 4. ABC curves of fish community in the Juma River in different seasons2.4 拒马河鱼类群落结构与环境因子的关系
DCA排序结果显示四个排序轴长度均小于3(表 5), 其中一、二轴长分别为2.84和2.28, 累积解释物种变化率为13.74%和22.72%, 物种和环境与排序轴相关性为94.14%、87.37%, 拒马河鱼类群落参数对环境因子的响应呈线性关系, 因此拒马河鱼类群落与环境因子分析适用于RDA分析。
表 5 拒马河鱼类群落DCA分析Table 5. Detrended correspondence analysis of fish community in the Juma River排序轴编号No. Axis 1 Axis 2 Axis 3 Axis 4 特征值Eigenvalues 0.3306 0.2159 0.1402 0.0736 排序轴梯度长度Gradient length 2.84 2.28 2.13 1.81 物种累计解释变量百分比
Cumulative percentage explained variation of species (%)13.74 22.72 28.54 31.60 物种和环境因子与排序轴相关系数
Species-environment correlations0.9414 0.8737 0.8688 0.8913 RDA分析中第一轴和第二轴为主成分轴(表 6), 第一轴解释变化率为21.73%, 前两轴累计解释变化率为29.03%, 物种和环境同前两轴相关关系分别为94.66%和81.29%。拒马河鱼类群落与环境因子RDA排序图结果(图 5)显示, 影响拒马河鱼类群落结构组成影响最大的环境变量为海拔(ASL), 对主要种类的解释量为20.60%, 是显著影响因子(蒙特卡洛检验F=8.8, P=0.002), 海拔与pH(蒙特卡洛检验F=0.2, P=0.024)同第一轴的相关性较强。海拔对尖头高原鳅Triplophysa Cuneicephala Shaw & Tchang、达里湖高原鳅、尖头鱥、拉氏鱥、马口鱼Opsariichthys bidens Günther、蛇Saurogobio dabryi Bleeker、棒花鱼等影响较大。黑鳍鳈、刺鳅、瓦氏黄颡鱼Pelteobagrus vachellii Richardson等对pH的变化呈负相关, 而兴凯鱊Acanthorhodeus chankaensis Dybowski对pH的变化呈正相关。水温(蒙特卡洛检验F=2.8, P=0.008)与第二轴相关性较强, 对鲫、高体鳑鲏和乌苏里黄颡鱼Pelteobagrus ussuriensis Dybowski等的影响较大。
表 6 拒马河鱼类群落RDA分析Table 6. Redundancy analysis of fish community in the Juma River排序轴编号No. Axis 1 Axis 2 Axis 3 Axis 4 特征值Eigenvalues 0.217 0.073 0.047 0.034 累计解释变化率
Cumulative percentage explained variation (%)21.73 29.03 33.69 37.09 物种和环境因子与排序轴相关系数
Species-environment correlations0.9466 0.8129 0.7315 0.5583 累计解释拟合变化率
Explained fitted variation (cumulative)50.19 67.05 77.82 85.67 ![]()
图 5 拒马河鱼类与环境因子的RDA图Tem. 水温; Dep. 水深; DO. 溶解氧; Chl.a. 叶绿素a; SD. 透明度; ASL. 海拔; FNU. 浊度; PH. 酸碱度; sp1. 鲫; sp2. 兴凯鱊; sp3. 中华鳑鲏; sp4. 高体鳑鲏; sp5. 尖头鱥; sp6. 拉氏鱥; sp7. 宽鳍鱲; sp8. 马口鱼; sp9. 麦穗鱼; sp10. 点纹银; sp11. 中间银; sp12. 兴隆山小鳔; sp13. 黑鳍鳈; sp14. 棒花; sp15. 棒花鱼;sp16. 蛇; sp17. 泥鳅; sp18. 北方泥鳅; sp19. 大鳞副泥鳅; sp20. 花斑花鳅; sp21. 北鳅; sp22. 塞丽高原鳅; sp23. 尖头高原鳅; sp24. 达里湖高原鳅; sp25. 刺鳅; sp26. 黄鳝; sp27. 子陵吻鰕虎鱼; sp28. 林氏吻鰕虎鱼; sp29. 波氏吻鰕虎鱼; sp30. 福岛吻鰕虎鱼; sp31. 小黄䱂鱼; sp32. 圆尾斗鱼; sp33. 黄颡鱼; sp34. 瓦氏黄颡鱼; sp35. 乌苏里黄颡鱼; sp36. 鲇; sp37. 青鳉Figure 5. Redundancy analysis of fish community and environmental factors in the Juma RiverTem. temperature; Dep. depth; DO. dissolved oxygen; Chl.a. chlorophyll a; SD. transparency; ASL. altitude; FNU. formazin unit; pH. potential of hydrogen sp1. Carassius auratus; sp2. Acheilognathus chankaensis; sp3. Rhodeus sinensis; sp4. Rhodeus ocellatus; sp5. Rhynchocypris oxycephalus; sp6. Rhynchocypris lagowskii; sp7. Zacco platypus; sp8. Opsariichthys bidens; sp9. Pseudorasbora parva; sp10. Squalidus wolterstorffi; sp11. Squalidus intermedius; sp12. Microphysogobio hsinglungshanensis; sp13. Sarcochilichthys nigripinnis; sp14. Gobio rivuloides; sp15. Abbottina rivularis; sp16. Saurogobio dabryi; sp17. Misgurnus anguillicaudatus; sp18. Misgurnus bipartitus; sp19. Misgurnus dabryanus; sp20. Cobitis melanoleuca; sp21. Lefua costata; sp22. Triplophysa sellaefer; sp23. Triplophysa cuneicephala; sp24. Triplophysa dalaica; sp25. Sinobdella sinensis; sp26. Monopterus albus; sp27. Rhinogobius similis; sp28. Rhinogobius lindbergi; sp29. Rhinogobius cliffordpopei; sp30. Rhinogobius fukushimai; sp31. Micropercops swinhonis; sp32. Macropodus ocellatus; sp33. Tachysurus fulvidraco; sp34. Pelteobagrus vachellii; sp35. Pelteobagrus ussuriensis; sp36. Silurus asotus; sp37. Oryzias sinensis3. 讨论
3.1 拒马河鱼类群落组成结构特征
拒马河位于太行山迎风区北部, 北连西北山间盆地和北京西山地区, 西部以山西省为邻, 南部与大清河南支山区相接[31], 根据李国良[32]对河北淡水鱼类地理区系的划分, 拒马河流域大部分属于冀西山地, 海拔高度由500 m增至1000 m以上, 鱼类资源以条鳅科(赛丽高原鳅、达里湖高原鳅)、花鳅科(北方泥鳅Misgurnus bipartitus Dybowski、泥鳅)和鲤科中的雅罗鱼亚科(尖头鱥、拉氏鱥、马口鱼、宽鳍鱲)、亚科(麦穗鱼、点纹银、中间银)、鲃亚科(多鳞铲颌鱼Onychostoma macrolepis Bleeker)鱼类为主。根据历史记载, 拒马河鱼类资源并不丰富, 李国良[33]和王所安[20]记录了9种, 杨文波等[16]在2008年对拒马河北京段调查中监测到24种, 袁立来等[6]在2019—2020年对拒马河北京段调查中监测到33种。然而, 针对整个拒马河流域的调查, 近几十年未见报道。本研究在2019—2021年6次对拒马河鱼类资源调查中共发现37种, 与王鸿媛等[34]的记录相比减少了多鳞铲颌鱼、花䱻Hemibarbus maculatus Bleeker、黄线薄鳅Leptobotia flavolineata Wang、东方薄鳅Leptobotia orierntalis Xu, Fang & Wang和花斑副沙鳅Parabotia fasciata Dabry & Thiersant等, 这些鱼类对生态环境的改变极为敏感, 仅分布于水质清澈、溶氧量高和无污染的狭窄水域。综合之前调查[5, 6, 16], 这几种鱼类很可能已经在拒马河流域消失。受地理环境因素和人类活动的影响, 拒马河鱼类分布具有明显的区域性, 海拔在500 m以上的上游区域(组Ⅰ), 渔获物以花鳅科和条鳅科鱼类为主, 包括北鳅Lefua costata Kessler、达里湖高原鳅、赛丽高原鳅、尖头高原鳅、花斑花鳅Cobitis melanoleuca Nichols和北方泥鳅等; 海拔在200m以上的中游区域(组Ⅱ), 水生态环境受人类影响相对较小, 如S3和S4, 渔获物以棒花鱼、中间银为主要组成; 海拔在200 m以下的下游区域(组Ⅲ), 位于旅游景区, 水生态环境受到人类影响较大, 如自S5至S15, 渔获物以黑鳍鳈、麦穗鱼、小黄䱂鱼、高体鳑鲏和泥鳅等喜栖息小河流水环境具有一定耐受力的鱼类为主。组Ⅰ的平均相似性为51.01%, 小于组Ⅱ的66.92%和组Ⅲ的61.00%, 体现了拒马河上游地区相较于中下游地区更具生境多样性。
2019—2021年拒马河优势种有鲫、麦穗鱼、黑鳍鳈和宽鳍鱲等, 按照初次性成熟小于2龄, 最大体长小于24 cm的鱼类划分为小型鱼类的标准[35], 拒马河鱼类群落优势种几乎都是小型鱼类。从食性上看, 拒马河鱼类以杂食性为主, 马口鱼、黄颡鱼和刺鳅等偏肉食性鱼类较少, 表明拒马河鱼类群落高位营养级少, 水生态环境更适合杂食性鱼类的生存。生态类型方面, 亲流性的鱼类物种数较多, 符合内陆河流鱼类组成特点, 但优势种中喜缓流及静水的鱼类较多, 原因可能是拒马河拦河坝的修建导致河流片段化, 部分河段水流速度减慢甚至形成小型静水湖泊, 破坏了亲流性鱼类的捕食、产卵场所同时也为喜静水的鱼类提供了更合适的栖息环境所导致[9, 36, 37], 在设计河坝时, 应优先考虑设计过鱼通道及保护可能因河坝建设而受到影响的鱼类, 降低因河坝的建设而导致的水文环境的改变对其造成的影响。在此次鱼类资源调查中, 麦穗鱼在所有月份都作为鱼类群落的优势种出现, 具有较高的生态优势度。麦穗鱼适应能力强、繁殖力高、食性广, 同拒马河土著鱼类构成了竞争关系, 且吞食鱼卵, 干扰产卵场, 对土著鱼类的生存构成极大威胁[38]。对于麦穗鱼多的河段, 应定期监控其种群动态, 必要情况下可采取人为措施控制其种群数量。
多样性指数(H')和均匀度指数(J')是群落结构稳定的重要评价指标[39], 物种丰富度越高, 个体数分布越均匀, 群落结构越稳定, 多样性指数和均匀度指数也就较大, 反之, 物种受到环境的胁迫, 群落结构不稳定, 多样性指数则低。在本研究中, 拒马河鱼类群落夏季的多样性指数高于春、秋两季, 这可能与夏季水温上升鱼类的活动频率增加有关。鱼类群落结构及物种多样性可以衡量水体健康情况, 水生态环境恶化会直接影响鱼类的物种多样性, 拒马河鱼类群落的Shannon-Wiener多样性指数(H')、Margalef丰富度指数(DMa)及Pielou均匀度指数(J')均表明拒马河的水环境受到了一定程度的污染[40]。
3.2 拒马河鱼类群落稳定性
Warwick[41]于1986年提出丰度生物量比较曲线用于判断鱼类群落稳定性。ABC曲线将生物量优势度曲线和丰度优势度曲线放置在同一坐标系中, 通过两条曲线的分布来分析鱼类群落的受干扰程度[42]。ABC曲线基于r选择和k选择的传统进化理论, 群落结构稳定时, 群落以k选择种类(生长慢, 性成熟晚的大个体种类)为主要组成, 生物量优势度曲线位于丰度优势度曲线之上。随着干扰的增加, k选择的物种逐渐减少而r选择的物种(生长快, 性成熟早的小个体种类)逐渐增加, 丰度生物量比较曲线随之发生改变, 群落受到中度干扰时, 生物量优势度曲线与丰度优势度曲线相交; 群落受到严重干扰时, 生物量优势度曲线位于丰度优势度曲线之下[43]。根据李胜法等[43]的研究标准, 拒马河鱼类群落整体只有2020年夏季结构稳定未受干扰, 其他月份鱼类群落结构受到中等程度的干扰, 但在拒马河不同河段受干扰程度存在差异, 海拔500 m以上的两个位点, 平均采集鱼类143.91尾, 显著高于海拔低于500 m位点的48.19尾, 并采集到数量可观的尖头鱥、拉氏鱥、赛丽高原鳅等对水质要求高的鱼类, 可看出该河段鱼类群落受扰动程度较小。
3.3 拒马河鱼类群落结构与环境因子关系
拒马河鱼类多样性指数中个体数、物种数、多样性指数、丰富度指数在不同季节间呈现显著性差异, 表明拒马河鱼类群落主要受非生物因子的影响。Kadye等[44]研究证明, 温度、海拔、pH、溶解氧和距河口距离等是影响河流鱼类群落结构的主要环境因子。王卓等[45]在研究汉江平川段鱼类群落结构与环境因子的关系中发现电导率(Cond)、5日生化需氧量(BOD5)、pH和硫酸浓度是影响该江段鱼类群落结构的主要环境因子。本文通过RDA分析得出海拔是影响拒马河鱼类群落结构的主要环境因子, 对达里湖高原鳅、尖头高原鳅和北鳅等鳅科鱼类影响较大。水温作用于鱼类的分布、生长、繁殖和迁移, 对拒马河鱼类群落结构的影响仅次于海拔, 自拒马河源头至龙安大桥水温逐渐上升, 鲫和高体鳑鲏等温水性鱼类数量也逐渐增加。兴凯鱊与pH的变化呈正相关, 而棒花鱼、刺鳅和瓦氏黄颡鱼等同pH的变化呈负相关, pH会对鱼的摄食和生长产生影响, 也会影响鱼的感官、代谢、呼吸等生理过程[46]。溶解氧与波氏吻鰕虎鱼Rhinogobius cliffordpopei Nichols、福岛吻鰕虎鱼等小型肉食性鱼类相关性较强, 符合肉食性鱼类耗氧高的特点。叶绿素a也是影响拒马河鱼类群落结构的重要环境因子, 叶绿素a与浮游植物的密度密切相关, 对小黄䱂鱼和棒花鱼等杂食性鱼类影响较大。
4. 结论
本研究系统地调查了拒马河鱼类资源, 共发现鱼类37种, 隶属于5目11科, 补充了拒马河上游区域的物种组成和地理分布等缺乏的基本问题, 为拒马河鱼类资源开发利用和资源保护提供了基础数据。基于等级聚类分析(Cluster)和非参数多变量排序(NMDS)分析发现, 拒马河鱼类分布在空间上具有明显的区域性, 上游以适应清澈流水环境的花鳅科和条鳅科鱼类为主; 中上游以棒花鱼、中间银为主; 中下游以黑鳍鳈、麦穗鱼、小黄䱂鱼和泥鳅等具有一定耐受力的鱼类为主, 这对拒马河生态修复和水利工程修建能提供理论依据。通过群落优势种和丰度生物量比较曲线分析发现, 拒马河以耐污性较强的小型鱼类为主, 主要为鲫、麦穗鱼、黑鳍鳈和宽鳍鱲等, 鱼类群落结构受到一定程度的干扰; 冗余分析显示, 海拔是影响拒马河鱼类分布的主要环境因子。拒马河是大清河水系的主要河流, 目前已有“北京市房山区拒马河水生野生动物自然保护区”, 建议进一步根据拒马河独特的水生态环境划定生态保护红线, 加强对该流域生物多样性的保护。
附表 S1 拒马河鱼类分布Appendix S1. Fish species distribution in Juma River物种
Species采样地点Sampling site 拒马河
源头刁江
汇紫荆关
大桥清凉
涧小丰
口桥别岸 琅琊
河天花
板北石
门西河
口九渡 六渡 穆家
口千河
口龙安
大桥鲤形目Cypriniformes 鲤科Cyprinidae 鲤亚科Cyprininae 鲫Carassius auratus + + + + + + + + + + + + + + + 鱊亚科AcheilognatSpnae 兴凯鱊Acanthorhodeus chankaensis + + 中华鳑鲏Rhodeus sinensis + + + 高体鳑鲏Rhodeus ocellatus + + + + + + + + + + 雅罗鱼亚科Leuciscinae 尖头鱥Rhynchocypris oxycephalus + + + + + + + + + 拉氏鱥Rhynchocypris lagowskii + + + + + + + 襁亚科Danioninae 宽鳍鱲Zacco platypus + + + + + + + + + + + + 马口鱼Opsariichthys bidens + + + + + + 亚科Gobioninae 麦穗鱼Pseudorasbora parva + + + + + + + + + + + + + + + 点纹银Squalidus wolterstorffi + + + + + + + + + + 中间银Squalidus intermedius + + + + + 兴隆山小鳔Microphysogobio hsinglungshanensis + + + + + + + + + 黑鳍鳈Sarcochilichthys nigripinnis + + + + + + + + + + + 棒花Gobio rivuloides + + + + + + + + + + 棒花鱼Abbottina rivularis + + + + + + + + + + + + + 蛇Saurogobio dabryi + + + + 花鳅科Cobitidae 泥鳅Misgurnus anguillicaudatus + + + + + + + + + + + + + + 北方泥鳅Misgurnus bipartitus + 大鳞副泥鳅Paramisgurnus dabryanus + + + + + + 花斑花鳅Cobitis melanoleuca + + + + 条鳅科Nemacheilidae 北鳅Lefua costata + 赛丽高原鳅Triplophysa sellaefer + + + + 尖头高原鳅Triplophysacuneicephala + 达里湖高原鳅Triplophysa dalaica + + + + 合鳃鱼目SymbrancSpformes 刺鳅科Mastacembelidae 刺鳅Sinobdella sinensis + + + + + + + 合鳃鱼科Symbranchidae 黄鳝Monopterus albus + 鲈形目Percoidei 鰕虎鱼科Gobiidae 子陵吻鰕虎鱼Rhinogobius giurinus + + + + 林氏吻鰕虎鱼Rhinogobius lindbergi + + + + + + + + + 波士吻鰕虎鱼Rhinogobius cliffordpopei + + + + + + + 福岛吻鰕虎鱼Rhinogobius fukushimai + + + + + + + 沙塘鳢科Odontobuidae 小黄䱂鱼Micropercops swinhonis + + + + + + + + + + + + + + + 丝足鲈科Osphronemidae 圆尾斗鱼Macropodus chinensis + + + 鲇形目Siluriformes 鲿科Bagridae 黄颡鱼Pelteobagrus fulvidraco + + + + + + + + + + + + + 瓦氏黄颡鱼Pelteobagrus vachellii + + + + 乌苏里黄颡鱼Pelteobagrus ussuriensis + + + + + 鲇科Siluridae 鲇Silurus asotus + + + + + + + 颌针鱼Beloniformes 青鳉科Adrianichthyidae 青鳉Oryzias sinensis + 注: “+”表示现场调查采集到样本Note: “+” represents collected fish species in the surveys -
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图 1 环境因子和鱼类特性对洄游行为及过鱼设施效果的影响关系示意图
Figure 1. Diagram of the influence of environmental factors and fish traits on migratory behavior and fishway effectiveness
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图 2 松新鱼道水流诱鱼设施实拍图
Figure 2. Photograph of the water flow-based fish attraction facility at Songxin Fishway
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图 3 湘河鱼道声驱鱼设施实拍图
Figure 3. Photograph of the sound-based fish repulsion facility at Xianghe Fishway
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图 4 马马崖一级水电站集运鱼系统光诱鱼设施实拍图
Figure 4. Photograph of the light-based fish attraction facility in the fish transport system at Mamaya I Hydropower Station
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图 5 果多水电站集运鱼系统电驱鱼设施实拍图
Figure 5. Photograph of the electricity-based fish repulsion facility in the fish transport system at Guoduo Hydropower Station
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表 1 近十年(2014—2024年)国内外关于鱼类对水流因子响应的研究进展
Table 1 Advances in domestic and overseas studies on fish responses to hydraulic factors in the past decade (2014—2024)
研究方法
Study method实验鱼种
Fish species具体表现
Specific performance文献来源
Reference1﹕2.5物理模型 鲢
Hypophthalmichthys nobilis鲢在流速为0.15—0.45 m/s, 紊动能为0.020—0.043m2/s2, 紊动耗散率为0.020—0.065 m2/s3和应变率为2—7/s的通过次数较多, 且紊动能和流速是影响竖缝式鱼道中鲢通过时间的关键水力参数 [31] 原型鱼道 短须裂腹鱼
Schizothorax wangchiachii平行补水时鱼道进口平均吸引效率为26.47%, 垂直补水时鱼道进口平均吸引效率为13.11%, 补水角度和流量显著影响进口鱼类吸引时间 [50] 1﹕10物理模型 草鱼
Ctenopharyngodonidella在3种鱼道入口和河道的角度(30°、45°和60°)条件下, 入口角度从30°增加到60°时, 草鱼的能量消耗范围从1.26—3.59×10−3 J增加到3.32—7.33×10−3 J, 且鱼头部偏转角度增加 [51] 物理模型 异齿裂腹鱼
Schizothorax oconnor在鱼道高流速、低紊动能区域易发生折返现象, 而折返后鱼类多选择低流速、高紊动能区域重新上溯 [52] 数值模拟 / 竖缝式鱼道模型中, 延长转弯段上下游长度的同时增设整流导板可有效引导主流居中, 维持主流稳定, 避免转弯段内产生大尺度回流区, 具有良好的上溯条件 [53] 数值模拟 / 竖缝式鱼道模型中, 转弯角度大于90°时, 主流冲击程度河回流区尺度较大。在增设整流导板后, 转弯角度α的逐渐增加, 整流较好的导板位置趋于稳定 [54] 原型鱼道 虹鳟
Oncorhynchus mykiss当池堰式鱼道坡度为10%时, 最大紊动能增加到0.22 m2/s2; 当坡度为4%且堰距较大时, 最大紊动能减小到0.05 m2/s2, 达到鱼在堰间池室中休息的最佳条件 [55] 物理模型 鳙
Aristichthys nobilis鳙幼鱼上溯游泳行为主要受紊动强度的影响, 其偏好紊动强度范围在5.25—8.40 cm/s [56] 数值模拟 齐口裂腹鱼
Schizothorax prenanti鱼道模型3个进口均处于坝下左岸最大概率预测洄游路线上 [57] 数值模拟 四大家鱼
Four major Chinese carps当流速处于0.24—0.70 m/s时, 四大家鱼数量与流速呈正线性相关; 当水深值处于2.4—3.9 m时, 四大家鱼数量与水深呈正线性相关关系; 紊动能在0—0.0012 kg·m2/s2和0.0015—0.0040 kg·m2/s2时, 四大家鱼数量与紊动能分别呈正相关和负相关关系 [58] 
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表 2 近十年(2014—2024年)国内外关于鱼类对声因子诱驱响应的研究进展
Table 2 Advances in domestic and overseas studies on fish attraction and repulsion responses to acoustic factors in the past decade (2014—2024)
鱼种
Fish species声音类型
Sound type趋音性
Phonotaxis具体表现
Specific performance实验类型
Experimental type文献来源
Reference黑鲷
Sparusmacrocephalus300和400 Hz正弦波交替音 正趋音性 正弦波交替音对黑鲷的反应和聚集具有明显的效果 室内实验 [69] 鲢
Hypophthalmichthys molitrix短吻鳄吼叫声
0—2000 Hz扫频音
打桩声
船舶噪声负趋音性
负趋音性
相对无趋音性
相对无趋音性打桩声和鸣笛声对鲢无显著驱赶效果, 0—2000 Hz扫频音和短吻鳄吼叫声有显著驱赶效果, 且吼叫声效果更好 室内实验 [70] 草鱼
Ctenopharyngodonidellus1000 Hz单频音
复杂音(鱼游动声、引擎声、短吻鳄吼叫声、打桩声和游艇声)相对无趋音性
负趋音性单频音对草鱼无显著驱赶效果, 而复杂音对草鱼有显著驱赶效果, 体现在相比单频音显著更高的反应次数、趋音速度和运动时间 室内实验 [71] 美洲拟鲽
Pseudopleuronectes americanus船舶噪声 负趋音性 暴露于船舶噪声环境中的美洲拟鲽幼鱼的捕食行为明显少于无噪声对照组 室内实验 [72] 草鱼
Ctenopharyngodonidellus500—3000 Hz
单频音
扬子鳄吼叫声均负趋音性 草鱼对扬子鳄吼叫声敏感程度远高于单频音, 平均反应次数高达(5.0±0.9)次, 且发生逃窜行为 室内实验 [79] 草鱼
Ctenopharyngodonidellus500—3000 Hz
单频音
摄食声相对无趋音性
正趋音性单频音对草鱼无显著诱驱效果, 而摄食声对草鱼有显著聚集作用, 体现在相比单频音显著更高的趋音游泳速度和逗留时间 室内实验 [80] 拉萨裸裂尻鱼
Schizopygopsis younghusbandi
双须叶须鱼
Ptychobarbus dipogon
斯氏高原鳅
Triplophysa stoliczkae扬子鳄吼叫声 均负趋音性 三种鱼对扬子鳄吼叫声均表现回避反应, 且拉萨裸裂尻鱼和双须叶须鱼对吼叫声更为敏感 野外实验 [81] 拉萨裸裂尻鱼
Schizopygopsis younghusbandi扬子鳄吼叫声
汽艇噪声
打桩噪声负趋音性
相对无趋音性
相对无趋音性拉萨裸裂尻鱼对扬子鳄吼叫声表现为负趋音性, 且声强增加, 负趋音性增强, 但其对汽艇噪声和打桩噪声的负趋音性不显著, 且不受声强影响 室内实验 [82] 大黄鱼
Larimichthyscrocea船舶噪声 负趋音性 当噪声声压级<60 dB时, 大黄鱼幼鱼负趋音性不强烈, 但声压级增大, 趋避行为强度增大, 依次表现出: 游泳速度加快、鱼与鱼之间及鱼与桶壁之间发生碰撞、瞬间反应无序和跳跃等行为 室内实验 [83] 裸腹叶须鱼
Ptychobarbus kaznakovi摄食声
河流噪声
1000 Hz单频音均正趋音性 摄食声引发裸腹叶须鱼显著的正趋声性, 效果优于1000 Hz单频音和河流噪声 室内实验 [84] 裸腹叶须鱼
Ptychobarbus kaznakovi500—3000 Hz
单频音
扬子鳄吼叫声均负趋音性 只有15%的裸腹叶须鱼对单频音有一次反应, 而裸腹叶须鱼对扬子鳄吼叫声反应更剧烈, 平均有8.4次反应 室内实验 [85] 真鱥
Phoxinus phoxinus106—212 Hz倍频带噪声
1556—3112 Hz倍频带噪声
150 Hz正弦波
2200 Hz正弦波均负趋音性 真鱥对4种声音都会受到惊吓, 会降低平均群体游泳速度, 且低频正弦波音对群体行为的影响最大 室内实验 [86] 大鳞吸口鱼
Moxostomaaureolum
淡水石口鱼
Aplondinotusgrunniens
大口黑鲈
Micropterus salmoides
虹鳟
Oncorhynchus mykiss涡轮机噪声 负趋音性
正和负趋音性
相对无趋音性
相对无趋音性大鳞吸口鱼对持续的涡轮机噪声表现负趋音性, 保持远离声源; 淡水石口鱼在中等音量下, 先表现出负趋音性后表现出正趋音性, 而大口黑鲈和虹鳟对涡轮机噪声表现相对无趋音性 室内实验 [87]
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表 3 近十年(2014—2024年)国内外关于鱼类对光因子诱驱响应的研究进展
Table 3 Advances in domestic and overseas studies on fish attraction and repulsion responses to light factors in the past decade (2014—2024)
鱼种
Fish species偏好光色
Preferred light color偏好光强
Preferred light intensity具体表现
Specific performance实验类型
Experimental type文献来源
Reference点带硬头鱼
Craterocephalusstercusmuscarum
西氏后鳍鲑
Retropinnasemoni/ 100—200 lx 点带硬头鱼和西氏后鳍鲑在昼夜光周期的调节下表现出显著的日间偏好。光照时间延长时, 它们的活动范围显著增加, 且克服迁徙障碍的概率也大幅增加 室内实验 [91] 大西洋鲑
Salmo salar/ / 大西洋鲑在日照时间逐渐变短的季节, 鲑鱼会启动长距离洄游, 以确保在适宜的环境中完成繁殖 野外实验 [92] 斑马鱼
Danio rerio绿光
紫光
红光/ 个体斑马鱼显著偏好绿光, 群体斑马鱼显著偏好紫光、红光和绿光, 个体群体均对黄光无趋光性 室内实验 [94] 拉萨裂腹鱼
Schizothorax waltoni绿光
蓝光/ 拉萨裂腹鱼在绿光和蓝光下表现为正趋光性, 在红光和黄光下表现为负趋光性 室内实验 [96] 秦岭细鳞鲑
Brachymystaxtsinlingensis绿光 10—25 lx 秦岭细鳞鲑稚鱼个体对3种光照强度(1—5、5—10和10—25 lx)和4种光照颜色(黄、红、绿和蓝)无明显趋避性, 而群体对强光和绿光有明显趋避性 室内实验 [99] 草鱼
Ctenopharyngodonidellus/ 34.62—52.45 lx 草鱼幼鱼在静水、0.1和0.2 m/s流水情况下, 光强期望值分别为52.45、34.62和37.86 lx 室内实验 [103] 鲢
Hypophthalmichthys molitrix红光
蓝光
绿光3.268—5.444 lx
0.033—10.511 lx
3.776—9.833 lx鲢的应激程度在红和绿光条件下随照度增加先增后减, 活跃性单调递增; 鲢的应激程度在蓝光条件下单调递增, 活跃性波动上升 室内实验 [104] 拉萨裸裂尻鱼
Schizopygopsis younghusbandi蓝光/深蓝光 / 在动水无底质条件下, 拉萨裸裂尻鱼均偏好于蓝光或深蓝光, 次亮光; 在静水有底质条件下主要偏好于暗光, 在无底质条件下偏好于次亮光 室内实验 [106] 斑马鱼
Danio rerio红光
绿光
紫光/ 斑马鱼对红光、绿光和紫光的喜好程度较高, 对黄光的喜好程度较低 室内实验 [107] 齐口裂腹鱼
Schizothoraxprenanti绿光
白光100—1000 lx
0—1000 lx在100—1000 lx绿光条件下, 齐口裂腹鱼的尝试次数显著高于10—100 lx绿光以及同照度白光, 其他照度会引起其逃逸频率或滞留时间的提升 室内实验 [108] 大麻哈鱼
Oncorhynchus keta
马苏大麻哈鱼
Oncorhynchusmasou/ 7.05 lx
11.83—31.7 lx在强光条件下, 大麻哈鱼呈负趋光性, 未发现正趋光性; 而马苏大麻哈鱼无负趋光性, 但正趋光性随光强增大会减少 室内实验 [109] 白鲟
Acipenser transmontanus绿光 / 鲟鱼在白天和夜间条件下都表现出积极的趋光性, 且绿光引起的吸引力最大 室内实验 [110] 中华倒刺鲃
Spinibarbus sinensis绿光 / 中华倒刺鲃幼鱼鱼群更倾向于绿光, 其群体协调性和凝聚力相比白光有明显提高 室内实验 [111]
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表 4 近十年(2014—2024年)国内外关于鱼类对电因子响应的研究进展
Table 4 Advances in domestic and overseas studies on fish responses to electrical factors in the past decade (2014—2024)
鱼种
Fish species电学参数
Electrical parameters具体表现
Specific performance实验类型
Experimental type文献来源
Reference海七鳃鳗
Petromyzon marinus
虹鳟
Oncorhynchus mykiss垂直布置: 3种脉冲电压(68.57、90.39和113.57 V)、4种电压梯度(0.43、0.68、0.72和0.94 V/cm)和4种脉冲宽度(1.6、8.2ms、41.8ms和300ms)交互组合
水平布置: 3种脉冲电压(16、20和40 V)、3种电压梯度(0.05—0.34、0.06—0.42和0.15—0.85 V/cm)、脉冲频率3 Hz和脉冲宽度2ms交互组合垂直布置和水平布置的脉冲直流电鱼栅均能有效引导海七鳃鳗, 但垂直布置的电鱼栅对虹鳟的阻拦效果显著优于水平布置 室内实验 [112] 鳙
Hypophthalmichthys nobilis
鲢
Hypophthalmichthys-molitrix脉冲电压0.91 V/cm 在芝加哥的伊利诺斯河, 拦鱼电栅用于防止入侵鲢鳙上溯进入密歇根湖, 尽管拦鱼电栅对两种鱼的阻拦效果良好, 但在自身洄游需求和环境水流刺激下, 鱼类会在拦鱼电栅附近停留 野外实验 [115] 鲤
Cyprinus carpio脉冲电压30 V、电压梯度0.2—0.4 V/cm、脉冲宽度0.3 ms和脉冲间隔时间10ms 通电后, 在电场的作用下, 鲤通过拦鱼电栅的次数明显减少, 且在刺激期布设电栅, 鲤鱼离开电屏障的时间比刺激前和刺激后更长 室内实验 [116] 鲤
Cyprinus carpio脉冲电压70 V、脉冲宽度0.4ms和脉冲间隔时间6ms 垂直电栅在三个阶段中对鲤的季节性迁徙都有极好的阻拦效果, 阻拦率在95.8%—98.6% 野外实验 [117] 虹鳟
Oncorhynchus mykiss脉冲电压30 V、电压梯度0.2—0.4 V/cm、脉冲宽度0.3 ms和脉冲间隔时间10ms 通电后, 拦鱼电栅使虹鳟通过的时间明显减少, 且远离电栅时间更长。在停止通电后, 虹鳟仍保持远离电栅 室内实验 [118] 大眼狮鲈
Sander vitreus3种脉冲电压(0、60和80 V)和4种脉冲宽度(0.3ms、0.5ms和0.8ms)交互组合 脉冲直流电不仅减少了大眼狮鲈的靠近, 还导致其偏转避开电屏障的几率增加, 这导致了近80%的逃逸率 室内实验 [120] 鲢
Hypophthalmichthys molitrix3种脉冲电压(80、110和140 V), 3种脉冲频率(6、8和10 Hz)和3种脉冲宽度(10ms、20ms和30ms)交互组合 影响拦鱼电栅对鲢阻拦效果由大到小的因素依次为脉冲电压、脉冲宽度、电极布置方式、脉冲频率, 在静水条件下, 拦鱼电栅对鲢的平均阻拦率在水流速度增至0.3 m/s时仍可达100%, 但进一步增至0.5 m/s时阻拦率有一定下降 室内实验 [122] 草鱼
Ctenopharyngodonide-llus4种脉冲电压(120、160、200和240 V/m)、4种脉冲频率(2、4、6和8 Hz)和4种脉冲宽度(10ms、12ms、14ms和16ms)交互组合 影响拦鱼电栅对草鱼阻拦效果由大到小的因素依次为脉冲电压、脉冲频率、脉冲宽度; 静水条件下脉冲电压160V/m、脉冲频率6 Hz和脉冲宽度16 ms的拦鱼电栅效果最好 室内实验 [123] 欧洲鳗鲡
Anguilla anguilla3种脉冲频率(2 Hz单脉冲、2 Hz双脉冲和10 Hz单脉冲) 相比双脉冲, 单脉冲需要更低的电场强度才能引起欧洲鳗鲡抽搐, 而电场强度对其反应无影响 室内实验 [124] 鲢
Hypophthalmichthys molitrix3种脉冲电压(80、110和140 V), 3种脉冲频率(6、8和10 Hz)和3种脉冲宽度(10ms、20ms和30ms)交互组合 影响拦鱼电栅对鲢阻拦效果和损伤率由大到小的因素依次为电极布置方式、脉冲电压、脉冲宽度和脉冲频率, 且阻拦效果和损伤率随流速增加无显著差异 室内实验 [125] 虹鳟
Oncorhynchus mykiss3种脉冲电压(51.4、69.0±0.8和91.4±0.8 V)、电压梯度0.00—0.45 V/cm、脉冲频率30 Hz和脉冲宽度0—0.7ms 当脉冲宽度较大时, 虹鳟鱼幼鱼和成年虹鳟鱼避开了电栅, 当脉冲宽度为0.7ms时, 通过电栅的虹鳟数量比脉冲宽度为0.1ms时少, 且未发现电压梯度对阻拦效果的影响 室内实验 [126] 鲤
Cyprinus carpio3种电极布置方式(垂直于水流方向且两排电极间距为80 cm、垂直于水流方向且两排电极间距为160 cm及与水流方向呈68°且两排电极间距为80 cm)、3种脉冲电压(80、110和140 V)、3种脉冲频率(6、8和10 Hz)和3种脉冲宽度(10、20和30 ms) 影响拦鱼电栅对鲤阻拦效果由大到小的因素依次为电极布置方式、脉冲宽度、脉冲频率和脉冲电压。垂直水流间距80 cm布置、脉冲电压140 V、脉冲频率10 Hz以及脉冲宽度10ms拦鱼效果最佳, 且随着脉冲宽度的增加, 平均受伤尾数相应增加, 随着脉冲电压的增加, 平均受伤尾数降低 室内实验 [127] 点纹似白鱼
Alburnoidesbipunctatus
欧洲鳗鲡
Anguilla Anguilla脉冲电压38 V、脉冲宽度0.3ms和脉冲间隔时间7ms 拦鱼电栅对点纹似白鱼和欧洲鳗鲡的保护效果较好, 对前者的保护率可达96%, 对后者的保护率也有86% 室内实验 [128]
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表 5 近十年(2014—2024年)国内外关于鱼类对气泡幕响应的研究进展
Table 5 Advances in domestic and overseas studies on fish responses to bubble curtains in the past decade (2014—2024)
鱼种
Fish species气泡幕结构
Bubble curtain structure具体表现
Specific performance实验类型
Experimental type文献来源
Reference鲤
Cyprinus carpio30 cm孔距和2.5 cm气泡幕; 5 cm孔距和2.5 cm孔径梯度泡幕; 5 cm孔距和2.5 cm孔径粗泡幕 梯度泡幕和粗泡幕可以减少鲤在上溯和下行两个方向通过气泡幕率几率达75%—85% 室内实验 [129] 虹鳟
Oncorhynchus mykiss4种孔距(1.0、2.0、3.0 和4.0 cm), 4种孔径(0.5、1.0、1.5 和2.0 mm)和3种气量(60、120 和180 L/min)交互组合 光照环境下, 孔径1.0 mm、孔距2.0 cm和气量120 L/min的气泡幕阻拦效果最好, 可达(96.32±3.99)%, 而黑暗条件气泡幕的阻拦率有所降低, 为(87.48±2.55)% 室内实验 [131] 光倒刺鲃
Spinibarbus hollandi3种孔距(1.0、2.5和
4.0 cm), 3种孔径(1.0、1.5 和2.0 mm)交互组合静水时, 9种组合产生的气泡幕对光倒刺鲃均有较好的阻拦效果, 达90%以上。流水环境提升了阻拦效果, 而闪光对光倒刺鲃有吸引作用 室内实验 [134] 鲤
Cyprinus carpio
鳙
Hypophthalmichthysnobilis
鲢
Hypophthalmichthys molitrix15 cm孔距和3 mm孔径 气泡幕对鲢、鳙和鲤均具有显著且相似的阻拦效果, 使其通过原路径的次数减少了73%—80% 室内实验 [135] 鲤
Cyprinus carpio5 cm孔距和2.5 cm孔径梯度泡幕 气泡幕对下行迁移的鲤幼鱼表现出较强的阻拦效果, 而对上溯成鱼的阻拦效果则相对较弱 野外实验 [136] 大泷六线鱼
Hexagrammosotakii2种孔径(1.0和2.0 mm), 2种孔距(2.5和5.0 cm)交互组合 4种组合产生的气泡幕均有明显的阻拦作用, 其中孔径1.0 mm与孔距2.5 cm组合的气泡幕阻拦效果最好, 平均阻拦率为53.2% 室内实验 [137] 异齿裂腹鱼
Schizothorax oconnori3种气量(60、90和
120 L/min)3种气量的气泡幕均有明显的阻拦效果, 气量60 L/min的气泡幕阻拦率最高, 为81.3%; 而其他气量的气泡幕附近存在鱼类停留现象 室内实验 [138] 异齿裂腹鱼
Schizothorax oconnori2种摆放角度(与水流方向呈90°和45°)下, 3种气量(15、30和45 L/min)交互组合 相比于静水条件, 流水条件下的气泡幕的影响距离更远, 垂直于水流方向布置效果更好, 且异齿裂腹鱼在尝试6次后会表现出适宜性 室内实验 [142] 鳙
Hypophthalmichthysnobilis
鲤
Cyprinus carpio
大口黑鲈
Micropterus salmoides30 cm孔距和2.5 cm
气泡幕声波与气泡幕结合对三种鱼相比单独布置气泡幕有更好的阻拦效果, 当和20—2000 Hz的循环声音结合, 可有效阻拦97%的入侵鳙 室内实验 [143] 大西洋鲑
Salmo salar1 cm孔距和1 mm孔径 在光线充足的白天, 气泡幕对大西洋鲑幼鱼的引导效率在实验室条件下达到95%, 自然环境中也能达到90%; 而在黑暗环境下, 效果显著降低 室内实验和野外实验 [144] 食蚊鱼
Gambusia affinis/ 日间条件下, 气泡幕显著增加了其两侧的鱼类数量, 但夜间气泡幕两侧的诱鱼效果并不显著 野外实验 [145] 大鳞大麻哈鱼
Oncorhynchus tshawytscha2.0 L/s的单位长度气泡速率 气泡幕与频闪灯和噪声发生器结合组成的生物声学鱼屏障, 有效引导了大鳞大麻哈鱼幼鱼避开低生存率的迁徙路线 野外实验 [146] 大泷六线鱼
Hexagrammosotakii3种气量(60、120和
180 L/min)3种气量的气泡幕对大泷六线鱼均有明显的阻拦效果, 且180 L/min气泡幕阻拦效果最好 室内实验 [147] 鲢
Hypophthalmichthys molitrix8种气量(10、20、30、40、60、80、100和
120 L/min)8种气量的气泡幕对鲢幼鱼都有阻拦效果, 其中, 20和30 L/min气泡幕阻拦效果最好, 气泡幕的气量与阻拦效果呈先上升后下降的趋势 室内实验 [150]
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