光照强度对大口黑鲈游泳协作能力的影响.pdf

doi:10.7541/2023.2023.0184光照强度对大口黑鲈游泳协作能力的影响吕义淞1,2 赵 建2 文彦慈2 杭晟煜2 倪伟强2 蔡海莺1 叶章颖2,3(1.浙江科技学院生物与化学工程学院,杭州 310023;2.浙江大学生物系统工程与食品科学学院,杭州 310058;3.浙江大学海洋研究院,舟山 316021)摘要:文章以大口黑鲈(Micropterus salmoides)为研究对象,研究了光照强度对大口黑鲈游泳能耗及游泳协作能力的影响,测定了4种不同光照强度(0、300500、22002500和33003900 lx)对大口黑鲈鱼群的游泳能耗、相对距离、相对位置分布和攻击次数的影响。研究发现:(1)当光照增强时,由于强光刺激和攻击行为的加剧显著制约了鱼群游泳协作能力的表达;(2)在光强22002500 lx时,随着光照强度的增大,鱼群游泳能耗显著增加(P蓝色红色绿色橙色黄色,且该品种对空间位置优先择取鱼缸的左右两侧18。当前光照对鱼类行为影响的研究主要集中在对养殖品种个体趋光性(光强和光色)的探究上,少有关注鱼类群体行为,尤其是其游泳协作能力。游泳能耗是鱼类日常能量支出中的主要部分,最高可达40%19,20,在很大程度上影响着养殖效益。鱼群内部的游泳协作可使群内个体相对于单独游泳时的游泳能耗降低(理论上或可高达70%21),避免不必要的能量损耗22,23。研究表明,环境因素直接影响着鱼类游泳协作能力。Hang等24发现在工业化水下噪声的影响下,大口黑鲈的群体结构较为松散,具体表现为相邻鱼之间的角度、距离增大;Paolo等25发现低氧状态时,为保证个体鱼能够尽可能多地从低氧水体中获取氧气,鱼群中个体之间的相对距离增大;Cooper等26发现当水温分布不均匀时,具有“偏好温度(Preferred temperatures)”的个体更倾向于停留在其偏爱的温度区域,而选择不跟随鱼群;流速则会导致鱼群的阵型配置发生改变,高流速下的鱼群更倾向于采取并排排列的阵型,而非低流速下的“钻石(Diamond-shaped)”阵型23;此外,Armstrong等27发现暴露在油污中会使鱼群中的个体平均移动速度降低及相邻个体之间的距离增大,从而削弱了鱼群的凝聚力(Cohesion)。然而,光照强度对鱼群游泳协作能力的影响还鲜有报道。大口黑鲈由于经济价值高,国内淡水养殖热门品种,被誉为继“四大家鱼”后的第五大家鱼。本研究以大口黑鲈为研究对象,探究不同光照强度对循环水养殖下鱼类游泳协作能力的影响,旨在为循环水养殖大口黑鲈光环境调控提供理论依据。1 1 材料与方法 1.11.1 实验材料大口黑鲈于2022年11月购自杭州唯康农业开发有限公司,总数为150尾,购买后置于本实验室规格为高120 cm直径100 cm的循环水圆形池内驯养15d。驯化期间每天上午10:00以通威商业颗粒饲料饱足投喂1次,投喂1h后立即清除残饵和粪便,日换水量约为驯化水体的25%。水温控制在(231)并保持24h不断曝气,水体溶氧水平为(6.20.5)mg/L,光周期为自然光照周期(12L12D)。在驯化完毕后选择个体健康且大小相近大口黑鲈体重(255)g,体长(152 cm)共66条作为实验对象。1.21.2 实验系统本实验系统主要由3部分组成:Brett-type游泳隧道28、摄像录制系统及照明系统。游泳隧道的主要作用是提供一个流速稳定可控的实验环境,摄像录制系统用于视频数据的录制、存储与分析,照明系统用于提供实验光源。游泳隧道(图 1)主要由转速可控的电机和内、外水槽(材料:透明亚克力)构成。电机通过频率控制器改变频率从而达到对其转速的精准控制,电机输出端为直径30 cm的螺旋桨,通过螺旋桨的转动在内水槽中产生稳定流速的非湍流水体29,水体温度保持在2224,流速为0.7 BL/s(体长/s)。内水槽(长180 cm宽80 cm高40 cm)开设长100 cm宽30 cm高30 cm的游泳槽,游泳槽内水体与实验对象直接接触。外水槽(长220 cm宽110 cm高40 cm)的主要作用保证内外水槽液面高度一致(不存在压强差),从而确保游泳槽内的流速恒定及整个Brett-type游泳隧道的稳定运转。摄像录制系统主要由相机(BASLER acA2040-电机频率控制器螺旋桨外水槽内水槽内水槽盖板挡流板1蜂窝板铁拦网挡流板2游泳槽图 1 Brett-type游泳隧道结构示意图Fig.1 Schematic diagram of the structure of the Brett-type swimming tunnel276水 生 生 物 学 报48 卷90 m,分辨率:2448680,帧率:60 fps)、刻录系统构成,相机架设于实验对象所在游泳槽的中央正上方,并通过Basler程序进行对焦和录制画面的参数调试。照明系统由红外补光灯和LED灯构成。红外补光灯(功率:90 W,长22.5 cm宽18.5 cm高15 cm。)的作用是给黑暗条件下的实验环境提供摄像机可见光源,红外补光灯架设在高度、角度可灵活调节并锁止的相机支架上。LED灯管(功率:035W可调,长118.5 cm宽2.1 cm高3.2 cm)通过底部固定、高度相同且角度可调节并锁止的灯管夹平行架设在实验对象所处游泳槽正上方,以保证游泳区域内光照相对匀称。光照强度检测通过光照强度测试仪(希玛AS823)在游泳槽注满水后由浅至深测量。1.31.3 实验方法实验开始前,先挑选一对体型大小接近的实验对象放入实验室内规格为长30 cm宽35 cm高45 cm的暂养水箱水温(231)中饥饿处理48h,以排空肠内残余消化物,避免特殊动力作用对实验对象耗氧率或游泳能力的影响30。具体操作步骤:(1)设置好实验灯光(每组鱼依次经历0、300500、22002500和33003900 lx的光强环境),并在禁食处理结束后将两条鱼用抄网轻轻放入游泳槽每组实验前将曝气24h、温度(231)溶氧水注入该游泳隧道。(2)给予实验对象30min休息时间,之后通过频率控制器调整输出频率,控制电机在2min内缓慢提升隧道内流速至0.7 BL/s,并在该流速下适应5min后通过置于游泳隧道正上方的摄像机拍摄记录(每个记录周期:记录3min+停止记录2min,每组实验对象共记录3个周期)。(3)在每种灯光实验结束后,将流速在2min内缓缓调节至0,并关闭灯光以供实验对象在黑暗中休息30min。(4)在休息结束后,改变灯光条件重复实验步骤(1)、(2)、(3)、(4)直至每组大口黑鲈经历完所有实验灯光。在本实验中,每组实验结束后重新更换游泳隧道中的水体,共计有33组实验对象完成本次实验。1.41.4 数据分析与处理游泳能耗量化指标鱼类在一般流速(0.05),但是总体上两组间的相对距离呈现出随着光强的增强而增大的趋势(图 5),同时在热度图中也可观察到当光强逐步增大时两鱼间距离的扩散变化过程(图 6)。大口黑鲈在光照强度不断增强的光环境下更倾向于采取前后排列的位置分布(图 5),且该位置分布的出现频率在各组间存在显著差异(P0.05)。热度图(图 6)反映了邻近鱼相对于焦点鱼的具体空间位置分布情况。2.32.3 不同光照强度下大口黑鲈鱼群攻击次数不同光照强度(分别为0、300500、22002500和33003900 lx)下大口黑鲈鱼群攻击次数分别为(2.390.629)、(3.790.76)、(6.731.36)和(7.061.62)次/180s,其中,33003900和22002500 lx下发生的攻击次数最高且与0组差异显著(P0.05)。总体呈现出攻击次数随光照的增强而逐步增大。3 3 讨论鱼群个体间相对距离和空间位置选择影响着鱼群游泳协作能力的表达,并继而决定了鱼群的游泳能耗32,38。鱼群个体间的距离和空间位置的选择由许多因素决定,如领导力、运动能力及饥饿程度35,39,40。本实验发现在黑暗条件下两鱼相对距离最短且很少采用前后排列(图 5),这可能是因为在短距离、左右排列的条件下两鱼可更好地利用彼此摆尾时产生的水动力学收益(Hydrodynamic be-nefits),从而节省游泳能耗22,41(图 4)。此外在黑暗条件下,两鱼间出现的攻击次数较少(图 7),在多数情况下采用安静、平稳的协作式游泳,这可能是为了减少自身受到捕食的风险并增加生存的机会42。330039002200250030050002.5cbaa2.01.51.00.50摆尾频率Tail beat frequency(Hz)n=33;x-SD光照强度Light intensity(lx)图 4 不同光照强度下大口黑鲈的摆尾频率Fig.4 The tail beat frequency of largemouth bass under differentlight intensity不同字母表示差异显著(P0.05);下同Values with different letters are significantly different(P0.05).The same applies below604020043210相对距离出现前后排列的频率出现前后排列的频率The frequency of front andrear arrangement occurrence(%)相对距离Relative distance(BL)33003900220025003005000n=33;x-SDdCcBbABaA光照强度Light intensity(lx)图 5 大口黑鲈在不同光照强度下的相对距离及出现前后排列的频率Fig.5 Relative distance and the frequency of front and rear ar-rangement occurrence of largemouth bass under different lightintensity278水 生 生 物 学 报48 卷研究发现,当光照增强时,两鱼表现为独立趋光探索、相互追逐的活动增强,协作游泳较少,具体体现在个体间的相对距离增大、更多地采取前后分布排列(图 5)及攻击现象加剧(图 7),同时伴随着高游泳能耗(图 4)。这一现象的出现可能是由于光强的增大导致了鱼群个体生理应激反应的增强,从而使得个体游泳时的活跃度及耗氧量增强43。类似现象也在其他研究中被报道,如:Schleuter等44发现欧亚鲈(Perca fluviatilis)在白天正午时的活跃度与傍晚相比提高了69.3%,正午时489 mg O2/(kgh)常规代谢率(Routine metabolic rate)相对于傍晚时220 mg O2/(kgh)提升了的1.22倍;而梅花鲈(Gym-nocephalus cernuus)正午时的活跃度相对傍晚时提高了61%,RMR则从176提升到289 mg O2/(kgh),约为64%;正午时大菱鲆(Scophthalmus maximus)的耗氧量248 mg O2/(kgh)显著高于早晨162 mg O2/(kgh)和傍晚171 mg O2/(kgh)45。值得一提的是,本研究中的最大光照强度33003900 lx对应的摆尾频率相较于22002500 lx有轻微下降但不明显(P0.05)。一般来说,对鱼类生长无明显影响的光照强度阈值为6002000 lx,当光照过强时,鱼类会展现出游泳疲劳和死亡率上升的现象46,47,比如海鲈(Di-centrarchus labrax)、露斯塔野鲮(Labeo rohita)和团头鲂(Megalobrama amblycephala)4749,这可能是本次实验中最高光强33003900 lx下摆尾频率相对22002500 lx组出现不显著下降的原因。事实上,光照的增强还会提高鱼群好斗性37,50,这可能也是制约鱼群游泳协作能力表达的重要因素。实验中观察到攻击现象的发生会导致两鱼间相对安静的平稳游泳状态被破坏,主要表现为个体对另一个体的撕咬、追逐及威胁(用嘴轻轻触碰另一个体的身体),进而导致两鱼长时间保持较远的相对距离,展现出彼此提防、警戒的现象,同时这也有可能是由于过强的光照会对群体内个体的视觉感知产生干扰,导致个体难以识别周围的伙伴或2.31.301.32.3X(BL)0.500.5Y(BL)000.0050.0100.0150.0200.0250.0302.31.301.32.3X(BL)0.500.5Y(BL)300500 lx00.0050.0100.0150.0200.0250.0302.31.301.32.3X(BL)0.500.5Y(BL)22002500 lx00.0050.0100.0150.0200.0250.0302.31.301.32.3X(BL)0.500.5Y(BL)33003900 lx00.0050.0100.0150.0200.0250.030 图 6 相对位置分布热度图Fig.6 The heatmap of relative position distribution12babaa108642033003900220025003005000光照强度Light intensity(lx)攻击次数Number of attack(180s)n=33;x-SD 图 7 不同光照强度下的攻击次数Fig.7 The number of attack under different light intensity2 期吕义淞等:光照强度对大口黑鲈游泳协作能力的影响279敌人而保持较远的相对距离51,52。此外,主动发起攻击的个体更经常地出现在群体前端,考虑到每条实验对象都经历了2d的禁食,因此主动发起攻击的个体可能更想通过占据前端获取优先进食的机会53,54。而被攻击的个体由于受到了频繁的攻击,更倾向于躲藏和减少活动,在本实验中观察到被攻击的个体一般会躲藏在攻击者的后端,且试图依赖水流方向拉开与攻击者间的距离,原因可能是为了减少暴露在攻击者的视野范围内,以降低再次被攻击的可能性。本研究表明,当光照强度较低时鱼群游泳协作能力较好,游泳能耗较低。而当光照增强时,鱼群游泳协作能力由于光强刺激、同类间攻击现象加剧而受到制约,因此,在实际生产中大口黑鲈养殖可更多地选择光强较弱(0500 lx)的养殖环境,以避免由于光照过强导致的鱼群游泳协作能力下降,降低鱼群游泳能耗及攻击现象的发生频次,减少生产养殖成本。参考文献:FAO.The State of World Fisheries and Aquaculture M.2020.1Zeng L Q,Fu C,Fu S J.The effects of temperature andfood availability on growth,flexibility in metabolic ratesand their relationships in juvenile common carp J.Com-parative Biochemistry and Physiology Part A:Molecular&Integrative Physiology,2018(217):26-34.2Christensen E A F,Svendsen M B S,Steffensen J F.Plasma osmolality and oxygen consumption of perchPerca fluviatilis in response to different salinities andtemperatures J.Journal of Fish 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