Undulatory fish swimming: from muscles to flow

Undulatory swimming is employed by many fish for routine swimming and extended sprints. In this biomechanical review, we address two questions: (i) how the fish's axial muscles power swimming; and (ii) how the fish's body and fins generate thrust. Fish have adapted the morphology of their axial musculature for high power output and efficiency. All but the superficial muscle fibres are arranged along curved trajectories, and the myomeres form nested cones. Two conflicting performance goals shape the fibre trajectories of the axial muscles. Maximum power output requires that all fibres contract uniformly. In a bending fish, uniform contraction in a single myomere can be ensured by curved fibre trajectories. However, uniform strain is only desirable if all muscle fibres have the same contractile properties. The fish needs several muscle‐fibre types that generate maximum power at different contraction speeds to ensure effective muscle power generation across a range of swimming speeds. Consequently, these different muscle‐fibre types are better served by non‐uniform contractions. High power output at a range of swimming speeds requires that muscle fibres with the same contractile properties contract uniformly. The ensuing helical fibre trajectories require cone‐shaped myomeres to reduce wasteful internal deformation of the entire muscle when it contracts. It can be shown that the cone‐shaped myomeres of fish can be explained by two design criteria: uniform contraction (uniform strain hypothesis) and minimal internal deformation (mechanical stability hypothesis). So far, only the latter hypothesis has found strong support. The contracting muscle causes the fish body to undulate. These body undulations interact with the surrounding water to generate thrust. The resulting flow behind the swimming fish forms vortex rings, whose arrangement reflects the fish's swimming performance. Anguilliform swimmers shed individual vortex rings during steady swimming. Carangiform swimmers shed a connected chain of vortex rings. The currently available sections through the total flow fields are often not an honest representation of the total momentum in the water – the wake of carangiform swimmers shows a net backward momentum without the fish accelerating – suggesting that our current picture of the generated flow is incomplete. To accelerate, undulatory swimmers decrease the angle of the vortex rings with the mean path of motion, which is consistent with an increased rate of backward momentum transfer. Carangiform swimmers also enlarge their vortex rings to accelerate and to swim at a higher speed, while eel, which are anguilliform swimmers, shed stronger vortex rings.     

流速对鲫游泳行为和能量消耗影响的研究

通过自制的鱼类游泳试验装置,研究了流速对鲫（Carassius auratus）游泳行为和能量消耗的影响。结果表明,鲫的摆尾频率、摆尾幅度随游泳速度变化有明显的规律。随着游泳速度的增加,鲫的摆尾频率与幅度都相应地增加。流速小于3倍体长/s与大于3倍体长/s,摆尾频率差异性显著（P〈0.05）,而摆尾幅度差异性不明显。鲫单位时间耗氧率随着流速的增加而显著增加,并且随着速度的增加,鲫用于游泳所消耗的能量占总能量消耗逐渐趋于稳定,运动净耗氧率最大为90%。水温（16±1）℃时,体长12～20cm鲫的相对极限流速为其（3.85±1.10）倍体长/s,绝对极限流速为（0.66±0.10）m/s;相同流速的温度环境中,随着体重的增加,耗氧量呈增加趋势。在相同的流速下,水温（10±1）℃时的游泳耗氧量小于（16±1）℃,而净耗氧率较大;温度（16±1）℃时,2.5倍体长/s的游泳速度有利于鲫生长代谢的能量积累。     

鲢幼鱼非疲劳贴网及姿态转换行为研究

流速是影响鱼类生存与繁衍栖息的重要因子。为了了解鱼类在中等流速区的游泳行为,探究其对水流的适应特征,利用鱼类游泳能力测定装置,以鲢(Hypophthalmichthys molitrix)幼鱼[体重(9.82±3.81) g,体长(8.56±1.11) cm)]为对象,采用递增流速法,分析了游泳过程中的非疲劳贴网行为与姿态转换行为。结果表明,在22℃水温条件下,鲢幼鱼平均相对临界游泳速度(critical swimming speed,U_(crit))相比其体长(body length,BL)为(6.00±0.93)BL/s;其非疲劳贴网速度(no-fatigue impingement speed,U_(imp))与临界游泳速度呈线性正相关,U_(crit)=1.03 U_(imp)+1.26 (R~2=0.86,P0.01);姿态转换速度(gait transition speed,U_(tran))与临界游泳速度呈线性正相关,U_(tran)=0.59 U_(crit)+1.55 (R~2=0.43,P0.01);非疲劳贴网速度与姿态转换速度呈线性正相关,U_(tran)=0.51U_(imp)+2.72 (R~2=0.41,P0.001)。在中等流速范围内(2～4 BL/s),实验鱼摆尾频率(tail beat frequency, TBF)和单次摆尾周期前进距离(stride length, SL)均随流速增加而增大。首次出现非疲劳贴网的流速为4.62 BL/s,流速增至5.08 BL/s时出现姿态转换行为。发生姿态转换后,TBF开始下降,而SL随流速增加快速增大。研究显示,鲢幼鱼非疲劳贴网行为对游泳能力和姿态转换速度有显著影响,且非疲劳贴网行为与姿态转换行为之间也存在相互影响。研究结果可为自然环境中鱼类生态行为研究、鱼类资源保护及渔业管理提供参考。     

A new device for monitoring the activity of freely swimming flatfish, Japanese flounder Paralichthys olivaceus

ABSTRACT:   The tail beat and activity behavior of four captive Japanese flounder Paralichthys olivaceus , were monitored with acceleration data-loggers while the fish swam in an aquarium. Depth, swimming speeds and two-axis acceleration data were collected continuously for approximately 20 h per fish. Simultaneously, the swimming behaviors of the fish were filmed at different angles. Using the specific characteristic of the acceleration profiles, in tandem with other types of data (e.g. speed and depth), four behavioral patterns could be distinguished: (i) 'active' swimming; (ii) burying patterns; (iii) 'inactive' gliding; and (iv) lying on the bottom. Tail beat frequency ranged from 1.65 ± 0.47 to 2.04 ± 0.25 Hz (mean ± SD; n  = 4). Using the relationship between tail beat frequency and swimming speed, the 'preferred' swimming speed of the fish was estimated to be between 0.6 and 1.2 body lengths (BL)/s. Additionally, fish rarely swam faster than 1.2 BL/s. This study shows that the acceleration data-loggers represent a useful and reliable system for accurately recording the tail beat of free-ranging fish and estimating flatfish behavior.     

异齿裂腹鱼游泳能力初探

为了摸清雅鲁藏布江特有种异齿裂腹鱼(Schizothorax oconnori)的游泳能力,以野生鱼种为实验对象,通过丹麦Loligo System公司的环形试验水槽测试了异齿裂腹鱼的临界游泳速度、突进游泳速度和持续游泳速度。结果显示,异齿裂腹鱼的临界游泳速度随体长的增大而增加,相对临界游泳速度随体长的增大而减小,其临界游泳速度与体长的关系为Y1=-39.369+13.23X-0.371X2+0.004X3(Y1是绝对临界游泳速度,X为体长)。突进游泳速度随体长的增加而近似呈线性递增趋势,而相对突进游泳速度随体长的增大而减小。在三个固定流速(60cm/s、80 cm/s、100 cm/s)下,初步确定60 cm/s为异齿裂腹鱼的持续游泳速度,80 cm/s、100 cm/s为耐久游泳速度。研究成果以期为青藏高原地区鱼类行为学的研究提供基础资料,为鱼道等过鱼设施的设计提供参考资料。     

Metabolic measurements and parameter estimations for bioenergetics modelling of Pacific Chub Mackerel Scomber japonicus

As a crucial step in developing a bioenergetics model for Pacific Chub Mackerel Scomber japonicus (hereafter chub mackerel), parameters related to metabolism, the largest dissipation term in bioenergetics modelling, were estimated. Swimming energetics and metabolic data for nine chub mackerel were collected at 14°C, a low temperature within the typical thermal range of this species, using variable‐speed swim‐tunnel respirometry. These new data were combined with previous speed‐dependent metabolic data at 18 and 24°C and single‐speed (1 fork length per second: FL/s) metabolic data at 15 and 20°C to estimate respiration parameters for model development. Based on the combined data, the optimal swimming speed (the swimming speed with the minimum cost of transport, U_opt) was 42.5 cm/s (1.5–3.0 FL/s or 2.1 ± 0.4 FL/s) and showed no significant dependence on temperature or fish size. The daily mass‐specific oxygen consumption rate (R, g O₂ g fish^?1 day^?1) was expressed as a function of fish mass (W), temperature (T) and swimming speed (U): R = 0.0103W^?0.490 e^(0.0457T) e^(0.0235U). Compared to other small pelagic fishes such as Pacific Herring Clupea harengus pallasii, Pacific Sardine Sardinops sagax and various anchovy species, chub mackerel respiration showed a lower dependence on fish mass, temperature and swimming speed, suggesting a greater swimming ability and lower sensitivity to environmental temperature variation.     

The effects of swimming exercise and dissolved oxygen on growth performance,fin condition and precocious maturation of early‐rearing Atlantic salmon Salmo salar

Swimming exercise, typically measured in body‐lengths per second (BL/s), and dissolved oxygen (DO), are important environmental variables in fish culture. While there is an obvious physiological association between these two parameters, their interaction has not been adequately studied in Atlantic salmon Salmo salar. Because exercise and DO are variables that can be easily manipulated in modern aquaculture systems, we sought to assess the impact of these parameters, alone and in combination, on the performance, health and welfare of juvenile Atlantic salmon. In our study, Atlantic salmon fry were stocked into 12 circular 0.5 m³ tanks in a flow‐through system and exposed to either high (1.5–2 BL/s) or low (<0.5 BL/s) swimming speeding and high (100% saturation) or low (70% saturation) DO while being raised from 10 g to approximately 350 g in weight. Throughout the study period, we assessed the impacts of exercise and DO concentration on growth, feed conversion, survival and fin condition. By study's end, both increased swimming speed and higher DO were independently associated with a statistically significant increase in growth performance (p < .05); however, no significant differences were noted in survival and feed conversion. Caudal fin damage was associated with low DO, while right pectoral fin damage was associated with higher swimming speed. Finally, precocious male sexual maturation was associated with low swimming speed. These results suggest that providing exercise and dissolved oxygen at saturation during Atlantic salmon early rearing can result in improved growth performance and a lower incidence of precocious parr.     

圆口铜鱼幼鱼可持续游泳能力及活动代谢研究

以圆口铜鱼(Coreius guichenoti)幼鱼为研究对象,通过自制的鱼类游泳实验装置,测定了4个温度(10,15,20和25℃)下圆口铜鱼幼鱼游泳速度达到临界游速过程中及运动疲劳后耗氧率的变化情况;在自然水温条件下,测定了5个不同流速下的可持续游泳时间,并通过摄像记录分析了不同游泳速度下的游泳行为。结果显示:圆口铜鱼幼鱼在运动疲劳前耗氧率随流速的增加显著上升(P0.05),在临界游速时达到峰值;运动疲劳后耗氧率逐渐下降,40~50 min内耗氧率恢复至低流速时的水平。自然水温(18.0±1.5)℃条件下幼鱼的可持续游泳时间随流速增加逐渐减小(P0.01),在1.31倍临界游速的固定流速下,平均可持续游泳时间达29 min,体现了较强的游泳耐力及无氧代谢能力。录像分析表明,摆尾频率(TBF)与游速的关系呈线性正相关(P0.001),且随着温度升高,TBF随流速增加的趋势越明显。     

水流对杂交鲟幼鱼游泳行为的影响

在26℃水温下,使用特制的鱼类游泳行为测定装置研究了杂交鲟(Huso duricusGeorgi♂×Acipenser schrenc-kiBrandt♀)幼鱼在0 m/s、0.1 m/s、0.3 m/s、0.5 m/s 4种流速条件下的游泳行为。结果显示:杂交鲟幼鱼随着流速的增加趋流率增大,且在90min内趋流率达到100%所需的时间明显缩短。实过过程中0.3 m/s和0.5 m/s两组摆尾频率都高于0.1 m/s和静水组,但在各个时段内,摆尾频率与趋流率均没有显著的相关。杂交鲟幼鱼的游泳状态明显受到所处流速的影响:0.1 m/s流速下以逆流前进为主,约占时间比例56.3%;0.3 m/s流速下以逆流静止为主,约占时间比例58.1%;0.5 m/s流速下以逆流后退为主,约占时间比例80.7%。在逆流前进和逆流静止两种游泳状态下,杂交鲟幼鱼的游泳速度和摆尾频率呈线性相关。但在逆流后退和顺流而下两种状态下,两者之间却没有显著的相关,且此两种状态下其游泳速度和摆尾频率随流速增加的变化趋势也不一致。     

不同运动状态下鳙幼鱼的游泳特性研究

为了探究鱼类反复运动疲劳后游泳能力和游泳行为的变化,以鳙(Aristichthys nobilis)幼鱼为研究对象,利用环形水槽实验装置,采用流速递增法,通过LoliTrack视频软件分析鱼类游泳行为,研究不同运动状态下鳙幼鱼的游泳特性变化情况。结果表明,鳙幼鱼疲劳前耗氧率随游泳速度的增大显著增加,且临界游泳速度恢复过程中最大耗氧率小于突进游泳速度恢复过程中最大耗氧率。视频分析显示,摆尾频率与游泳速度呈线性正相关关系。同时发现,在2组反复疲劳运动状态下,连续进行2次临界游泳速度测定过程中,鳙幼鱼频繁使用爆发-滑行游泳行为,而在经过突进游泳速度测试后转入临界游泳速度测定过程中,实验鱼并未使用这一游泳行为。经过一次临界游泳速度测试后,第二次临界游泳速度显著大于突进游泳速度测试后的临界游泳速度(P0.05),第一次突进游泳速度与第二次突进游泳速度无显著差异(P0.05)。无氧运动消耗导致鳙幼鱼有氧运动能力显著下降,而有氧运动消耗过程对无氧运动能力无显著影响。     

人工繁殖圆口铜鱼幼鱼游泳能力与游泳行为研究

利用鱼类游泳能力测定装置,以人工繁殖的圆口铜鱼(Coreius guichenoti)幼鱼为实验对象,体重(3.05±0.99)g,体长(5.94±0.66)cm,采用递增流速法研究其游泳能力与游泳行为。结果表明,在(20±1)℃水温条件下,圆口铜鱼平均相对临界游泳速度(critical swimming speed,U_crit)为(8.41±1.56)BL/s;其步态转换速度(gait transition speed,U_chg)与临界游泳速度呈线性正相关:U_chg=0.63 U_crit+0.21(P<0.01,R 2=0.86);摆尾频率(tail beat frequency,TBF)与进口流速(inlet velocity,U_in)呈线性正相关:TBF=0.48 U_in+2.53(P<0.01,R 2=0.95);进口流速(inlet velocity,U_in)与步长(step length,SL)呈线性正相关:SL=0.11 U_in+0.41(P<0.01,R 2=0.99)。实验鱼的摆尾幅度(tail beat amplitude,TBA)、冲刺次数、各进口流速下的摆尾时间百分比及稳定摆尾与非稳定摆尾比例,均随进口流速改变而变化。随流速增大,摆尾幅度呈现先增大、再减小、最后又增大的趋势;冲刺次数也是先增加、随后逐渐下降;摆尾时间百分比最初是快速增大,随后基本保持不变,流速增至8 BL/s高流速时,再次随流速增加而快速增大。当流速与临界游速比值(U/U max)为0.38时,实验鱼出现非稳定摆尾行为;比值为0.58时,实验鱼稳定摆尾行为与非稳定摆尾行为比例为1∶1;比值增至0.78时,稳定摆尾行为消失。人工繁殖的圆口铜鱼游泳能力较强,在不同流速下,通过改变游泳行为以保持更长的游泳时间及距离。研究结果可为以圆口铜鱼为过鱼对象的鱼道建设以及养殖流速优化提供参考。     

Temperature and fatigue effect on the maximum swimming speed of jack mackerel Trachurus japonicus

Swimming performance of jack mackerel Trachurus japonicus (18.2 ± 0.8 cm fork length (FL), n = 185) was examined in a flume tank by measuring the stride length at low and high tail beat frequencies with electromyogram monitoring and a muscle twitch experiment. Stride length was analyzed by monitoring the tail beat frequency according to the swimming speed at different temperatures of 10, 15 and 22 °C. In the electromyographic observations, the initiation of ordinary muscle activity occurred between 71.4 and 99.6 cm/s, that is 3.7 to 5.3 FL/s, when the tail beat frequency was over 6 Hz. The swimming speeds increased rectilinearly with the tail beat frequency at each water temperature both for the low and high tail beat frequency. Lower stride length was observed at the lowest temperature (10 °C) tested. The forced swimming exercise significantly affected the muscle contraction time to become longer than the control fish, which indicated a reduction of the maximum swimming speed performance.     

Use of electromyogram telemetry to assess the behavioural and energetic responses of rainbow trout, Oncorhynchus mykiss (Walbaum) to transportation stress

Behavioural and energetic responses of domesticated rainbow trout Oncorhynchus mykiss (Walbaum) (mean fork length=440±45 mm) to a brief transportation episode were investigated. Fish implanted with radio transmitters measuring muscle activity (electromyogram; EMGi) were transported in a standard commercial shipping tank for 50 min by truck, and then allowed to recuperate for 48 h in stationary culture tanks. The EMGi telemetry data indicated that vigorous swimming activity occurred during transportation. Telemetry recordings also indicated that the fish's swimming activity returned to baseline levels within the 48 h period after transport. However, even beyond the 48 h resting period, the swimming performance (measured as critical speed and endurance) of transported fish was still impaired relative to non‐transported controls (P<0.05). Respirometry measurements of fish taken after transportation indicated that oxygen consumption (V_o2) was significantly elevated. The rise in V_o2 of post‐transport fish could be attributed to handling procedures, as well as the intense swimming behaviour observed during transportation. Therefore, the behavioural responses of fish during transportation produced physiological consequences that persisted long after the transportation event. This study demonstrates the potential for utilizing behavioural measures, in concert with biotelemetry technologies, as tools to assess the impacts of routine aquacultural procedures on the health and welfare of captive fish.     

小头鲔幼鱼陆基循环水养殖日周期性运动规律研究

为实现小头鲔(Euthynnus affinis)陆基循环水养殖,优化养殖设施和完善养殖技术参数,开展了陆基循环水驯化养殖条件下,小头鲔口径与全长的关系、非摄食期间幼鱼运动速度、摄食期间幼鱼运动速度、幼鱼摄食深度与时间的关系及非摄食期间幼鱼水层分布等一系列研究。结果显示:小头鲔幼鱼45°和90°口径均随着全长的增长而增大,呈线性关系;在非摄食状态下,小头鲔幼鱼主要做巡航游泳运动;摄食阶段,初始发现食物时,小头鲔幼鱼做爆发性游泳运动,游泳速度与摄食游泳速度和摄食后游泳速度差异显著(P<0.05);捕食时和捕食后,小头鲔主要做巡航游泳运动,两者间差异不显著(P>0.05)。在摄食状态下,小头鲔幼鱼的游泳深度会随食物的深浅变化而变化;而在非摄食状态下,小头鲔幼鱼的游泳深度变化与时间有一定的关系。本研究为小头鲔陆基养殖、繁育等提供基础数据,也为后续开展南海金枪鱼陆基驯化养殖及人工繁育提供参考。     

小头鲔幼鱼陆基循环水养殖日周期性运动规律研究

为实现小头鲔(Euthynnus affinis)陆基循环水养殖,优化养殖设施和完善养殖技术参数,开展了陆基循环水驯化养殖条件下,小头鲔口径与全长的关系、非摄食期间幼鱼运动速度、摄食期间幼鱼运动速度、幼鱼摄食深度与时间的关系及非摄食期间幼鱼水层分布等一系列研究。结果显示:小头鲔幼鱼45°和90°口径均随着全长的增长而增大,呈线性关系;在非摄食状态下,小头鲔幼鱼主要做巡航游泳运动;摄食阶段,初始发现食物时,小头鲔幼鱼做爆发性游泳运动,游泳速度与摄食游泳速度和摄食后游泳速度差异显著(P<0.05);捕食时和捕食后,小头鲔主要做巡航游泳运动,两者间差异不显著(P>0.05)。在摄食状态下,小头鲔幼鱼的游泳深度会随食物的深浅变化而变化;而在非摄食状态下,小头鲔幼鱼的游泳深度变化与时间有一定的关系。本研究为小头鲔陆基养殖、繁育等提供基础数据,也为后续开展南海金枪鱼陆基驯化养殖及人工繁育提供参考。     

The effect of demand feeding on swimming speed and feeding responses in Atlantic salmon Salmo salar L., gilthead sea bream Sparus aurata L. and European sea bass Dicentrarchus labrax L. in sea cages

Abstract Using underwater cameras, data were collected on the feeding behaviour and swimming speeds of Atlantic salmon Salmo salar L., gilthead sea bream Sparus aurata L. and European sea bass Dicentrarchus labrax L. in sea cages. Comparisons were made between the behaviours of fish fed on demand using interactive feedback systems and those of fish fed under the standard feeding practice of each farm (control). In all three species, swimming speeds were similar before feeding , but they were significantly higher in the control regimes during feeding. When fed on demand, sea bass had reduced swimming speed just before and during feeding compared with that observed during the non‐feeding periods. Higher proportions of feeding fish were observed in the control regime cages than in fish fed on demand for all three species, indicating a greater feeding intensity during meals in the control regimes. This was further supported by observations of an increase in the density of sea bass in the upper water in the control cages during feeding. The results suggest decreased levels of competition between the on demand‐fed fish during feeding, which might be hypothesized to lead to improved growth and production efficiency in aquaculture.     

5种鱼类标志对草鱼临界游泳速度的影响

采用被动整合雷达标志法(PIT)、切鳍标记、荧光标记、超声波标志和T型标志这5种标志方法对草鱼(Ctenopharyngodon idellus)进行标记后,测定其临界游泳速度,研究5种不同标记对草鱼游泳能力的影响。将试验草鱼按不同体长分为3组:15~18 cm,18~21 cm,21~25 cm,每个体长组均设置对照组,并分别进行以上5种鱼类标志,测量标志后草鱼的绝对临界游泳速度和相对临界游泳速度。采用SPSS17.0统计软件进行数据的分析比较。研究结果表明,随着体长增加,草鱼的绝对临界游泳速度增大,相对临界游泳速度减小;超声波标志对3个体长组草鱼的临界游泳速度均有极显著影响(P0.01),标志后草鱼绝对临界游泳速度分别下降18.72%、16.40%、23.15%,相对临界游泳速度分别下降18.95%、17.78%、21.86%;T型标志对15~18 cm体长组草鱼的临界游泳速度有极显著性影响(P0.01),标志后草鱼绝对临界游泳速度下降8.35%,相对临界游泳速度下降9.30%,对体长大于18 cm的草鱼无显著性影响;PIT标志、切鳍标记和荧光标记对这3个体长组草鱼的临界游泳速度均无显著性影响(P0.05)。     

北盘江2种典型增殖放流鱼类突进游速研究

长臀（Cranoglanis bouderius）和白甲鱼（Onychostoma sima）是目前北盘江主要放流鱼类,但这2种鱼的放流效果差异较大。由于鱼类的突进游速在一定程度上影响增殖放流的效果,故试验采用自制测试装置,用递增流速法测试了2种鱼的突进游速。结果显示,白甲鱼的突进游速大于长臀,且2种鱼的突进游速均随体长增加而近似线性增大;而相对突进游速则随着体长的增加而近似线性减小。研究结果可为北盘江鱼类增殖放流和放流效果评估,以及日后北盘江的拦鱼、诱鱼、集鱼船等鱼类资源保护措施的实施提供参考。     

Ontogenetic behaviour and swimming ability of the endangered eastern freshwater cod,Maccullochella ikei,with notes on growth and development

Abstract – Understanding the larval ecology of individual fish species is fundamental in ensuring their long‐term conservation. The endangered eastern freshwater cod, Maccullochella ikei, is endemic to the Clarence and Richmond River systems of north‐eastern New South Wales, Australia. Little is known about the behaviour of larval M. ikei in the wild, particularly before and after the swim‐up stage, and following dispersal from the nest site. The aims of this study were to quantify the swimming ability, depth selection, light preference and substrate selection of hatchling to day‐30 M. ikei under controlled laboratory conditions, and to describe its growth and development over the same period. Maccullochella ikei larvae grew constantly but not consistently during the experiment. Exogenous feeding commenced around day 12, prior to the full exhaustion of the yolk. Maximal swimming ability improved daily, but maximum swimming speed declined significantly between days 12 and 13 and remained low. Maccullochella ikei larvae were initially photonegative but were positively phototactic by day 10. Depth selection was for the benthos until day 8, beyond which time larvae dispersed to all depths when released. Substrate selection was for sand in younger larvae but changed to upstream substrates as the experiment progressed. The results of the current study suggest that the period between day 10 and day 20 is critical in the early ontogeny of M. ikei, when it switched phototrophic behaviour, transitioned from endogenous to exogenous feeding and experienced a decline in swimming ability.     

Hemoglobin level,metabolic rate,opercular abduction rate and swimming efficiency in female triploid brook trout (Salvelinus fontinalis)

Metabolic (oxygen consumption) rate, opercular abduction rate and tail beat frequency were determined in two strains of diploid and triploid female brook trout (Salvelinus fontinalis) while these fish swam at 0.37±0.02 body lengths per sec in a Blazka respirometer. Total blood hemoglobin level was also measured and opercular condition examined. Total blood hemoglobin levels in diploids and triploids were equal. The opercular abduction rate was the same in diploids and triploids (regardless of whether triploid opercular condition was good or poor) yet triploids had a lower oxygen consumption rate than diploids, indicating that triploids take up less oxygen than diploids per opercular cycle. Tail beat frequency, an indicator of swimming effort, was the same in diploids and triploids, suggesting that triploids require less oxygen than diploids for a similar swimming effort.