Activities of mesencephalic nucleus neurons during fictive swimming of the carp Cyprinus carpio

ABSTRACT:   Fish swim by undulatory contractions of the axial trunk musculature. It has been presumed that a descending signal from the brainstem activates central pattern generators in the spinal cord to make the swimming rhythm. In the carp the electrical or chemical stimulation of a mesencephalic nucleus, the nucleus fasciculi longitudinalis medialis (Nflm), could initiate swimming. However, it has not been established what signals originate from the Nflm in order to make the fish swim. The activity of the Nflm neurons was therefore examined electrophysiologically during fictive swimming in paralyzed carp. Three types of neuronal activities closely related to the swimming rhythms were obtained. The first type was a continuous tonic firing throughout an episode of fictive swimming. Neurons involved in this tonic activity may project to the spinal cord and contribute to the activation of spinal neurons to initiate fish swimming. The second type was accompanied by continual phasic firings occurring in rhythm with the activity of spinal motoneurons. Supposing that the swimming rhythm originates in the spinal cord, the synchronous activity in the brain neurons may suggest that in the nucleus there is a relay neuron conveying the rhythm information from the cord to other neurons there or in the brain. The third type exhibited reduced firing rates during fictive swimming. It is possible that the neurons engaged in this activity may be inhibitory and suppress the activities of other neurons in the nucleus and spinal cord during rest or during decelerating swimming.     

Recording cerebellar neuron activities in swimming goldfish

ABSTRACT:   Neuronal activities were investigated in the cerebellum of immobilized and swimming goldfish Carassius auratus . Extracellularly recorded neural activities of the cerebellum in immobilized goldfish were characterized and classified into five types. Based on the waveforms and recording depths, these five neural activity types were estimated to originate from three identified classes of cerebellar neurons: Purkinje cells, eurydendroid cells, and granule cells. Chronic recording of cerebellar neuron activities in unrestrained goldfish was performed for more than 100 h. During the chronic recordings, a submersible amplifier attached to the goldfish head, and a multielectrode array developed for the present study were used. Neuronal activities in the cerebellum of free-swimming fish could also be classified into five types as in the immobilized condition. Firing patterns of two neurons identified as Purkinje cells and eurydendroid cells were analyzed during turning movements of the goldfish. The firing patterns of these neurons changed in relation to turning movements. Although some improvements are required, the chronic recording method developed in the present study can be applied to further investigations concerning the direct relationship between brain neural activities and certain behavior.     

Activity and energetics of free-swimming fish: insights from electromyogram telemetry

Electromyogram (EMG) telemetry studies that involve remotely monitoring the locomotory activity and energetics of fish are contributing important information to the conservation and management of fisheries resources. Here, we outline the development of this rapidly evolving field and formulate the studies conducted that utilize this technology. To date, more than 60 studies have been conducted using EMG telemetry that spans 18 species. Several general trends were observed in the methodology of the studies that we have highlighted as standards that should be adopted associated with transmitter customization, electrode placement and surgical technique. Although numerous studies have been methodological, there are still some deficiencies in our basic understanding of issues such as the need for individual calibration and the method of reporting or transforming data. Increasingly, this technology is being applied to address issues in conservation, management and aquaculture production. At present, the technology has been most frequently applied to the study of animal activity or energetics and to migration. Several recent studies have also focused on addressing more basic questions in ecological and evolutionary biology (e.g. parental care dynamics) similar to the large body of literature that has been collected for other taxa (e.g. marine mammals, birds), using activity telemetry. Collectively, studies conducted using EMG telemetry have contributed important information on free‐swimming fish that was previously difficult to obtain. EMG telemetry is particularly effective for examining behaviour at temporal and spatial scales that are difficult using other techniques. The development of an ultrasonic transmitter based on the same proven principles as those used in the current radio transmitter technology will permit studies in other environments (i.e. marine, brackish, deep water) and on different species of fish. We encourage the continued development and refinement of devices for monitoring the activity and energetics of free‐swimming fish, and also encourage researchers to consider EMG telemetry as a tool for addressing questions that are not effectively answered with other techniques.     

Muscle activity as a key indicator of welfare in farmed European sea bass (Dicentrarchus labrax L. 1758)

Groups of adult sea bass were reared at either low (10 kg m^?3) or high (50 kg m^?3) stocking densities respectively for 84 and 116 days. To monitor the red muscle activity, about 20 fish from both densities were surgically implanted with EMG (Electromyograms) radio transmitters, after EMG calibration during exhaustive swimming exercise (U_crit test). Blood samples and morphometric measurements were also taken. EMG showed that the muscle activity of fish reared at 50 kg m^?3 was on average twofold higher than fish kept at lower density. Cortisol was significantly more elevated at higher density and haemoglobin, haematocrit and RBCC (red blood cells count) showed the same trend, while lysozyme decreased. Patterns for glucose and lactate were less clear. The results showed that the contemporary use of functional (EMG) and physiological (haematological and biochemical) profiles could give a more comprehensive view of the fish status validating the diagnosis of fish stress induced by culture practices.     

Stunning of farmed African catfish (Clarias gariepinus) using a captive needle pistol; assessment of welfare aspects

The objective of this study was to assess whether high‐pressure injection of air into the brain of African catfish (Clarias gariepinus) could render the animal unconscious and insensible immediately and permanently. In the study, 48 African catfish with a live weight of 900–1900 g were restrained and equipped with EEG and ECG electrodes and then stunned. The catfish were stunned mechanically using a captive needle pistol. The pressure to shoot the needle was 8 bar and that to inject the air was 3 bar for 1.5 s. The catfish behaviour was observed during and after stunning. τ and δ waves and spikes, which precede a stoppage in brain activity as measured on the EEG, were used as indices for the measurement of immediate induction of unconsciousness and insensibility In 23 of 42 fish, an iso‐electric line was observed after an average of 13.4 s, while in the remaining fish the τ and δ waves and spikes remained on the EEG during the recording period. In all cases, the ECG showed an irregular heart rate with fibrillation and extra systolae. Moreover, the configuration showed ischaemia. Before the captive needle stunning, free‐swimming fish (n=7) explored the tank for an average of 21±12 s before lying down at the bottom. After stunning, they showed clonic uncoordinated swimming movements. The movements stopped after an average of 38±50 s. In another group (n=7) that was stunned and subsequently placed in ice water, clonic cramps were observed in two out of seven animals. When taking into account the number of animals with a reliable EEG (n=42) and using 95% confidence intervals, it was concluded that at least 93% of the catfish were effectively stunned using a correctly positioned captive needle pistol. Furthermore, it is recommended to immobilize the stunned fish by chilling, as the post‐stun clonic cramps may hinder gutting and filleting.     

Aerobic muscle function during steady swimming in fish

Axial swimming in fish varies across a range of body forms and swimming modes. Swimming by eels, tunas, mackerels, scup, rainbow trout and bass span this range from high curvature anguilliform swimmers to rigid body thunniform swimmers. Recent work on these and other species has elucidated an impressive array of solutions to the problem of how to use the red (aerobic, slow‐twitch) muscle to power steady or sustained swimming. This review will use a comparative approach to understand the generalities of aerobic muscle function during steady swimming in fish and determine possible rules for the relationships between muscle contractile kinetics, in vivo muscle activity and power output during swimming. Beyond an exploration of the diversity in muscle activity and swimming kinematics, I suggest that analysis of the molecular basis for longitudinal variations in muscle function is needed to complement morphological and physiological research on fish muscle. This will permit both a general understanding of the integrative function of the fish myotome and, perhaps, predictive tools for muscle activity and swimming performance in fish.     

Cardiovascular responses to acute handling stress in the Antarctic fish Trematomus bernacchii are not mediated by circulatory catecholamines

Cannulated specimens of the benthic Antarctic teleost Trematomus bernacchii were subjected to a five min period of handling stress, following which blood pressures were measured and then blood samples taken for measurement of plasma catecholamine concentrations. There were two control groups of fish, one group of undisturbed fish at Scott Base, Antarctica and the other group of undisturbed fish measured in Christchurch.Plasma catecholamine concentrations were not changed following 5 min of acute handling stress. However, heart rate, ventral aortic blood pressure and haematocrit were elevated following handling, suggesting increased sumpathetic nerve activity. The low plasma catecholamine concentrations measured in all three groups of fish are similar to those found in other species of teleost at rest.The resting heart rates found in this series of experiments were approximately double those recorded in previous work on this species and this difference may be correlated with the length of time that the fish are held in captivity. The fish transported to Christchurch also had higher mean cell haemoglobin concentrations (MCHC) and lower plasma chloride concentrations.     

Movements of Atlantic salmon migrating upstream through a fish-pass complex in Scotland

Abstract  – Movements of adult Atlantic salmon were tracked through a series of four fish passes and an impoundment on the River Conon system, Northern Scotland. Proportions of fish passing individual obstructions ranged from 63 to 100%. The cumulative effect was that only 4 of the 54 tagged fish reached the spawning areas. The fish were delayed for 1–41 days at a pool-and-overfall ladder and 1–52 days at a Borland fish lift. The fish swam through a 10 km long reservoir at 0.21–1.16 km·h⁻¹. A total of 13 fish negotiated a 2.5 km long, 3 m diameter diversion tunnel through a mountain to their home river. High levels of electromyogram (EMG) activity were recorded during ascent of a pool-and-overfall fish ladder, indicating that high energy demanding burst swimming was required.     

Effects of anesthesia and surgery on U crit performance and MO2 in chum salmon, Oncorhynchus keta

Telemetry is a useful technique for elucidating salmon behavior, but the recovery periods before fish can be safely released after the attachment of telemetry devices have not yet been established. Reported recovery times vary widely, from 2 h to 13 days. We examined how anesthesia and surgery to attach external electromyogram (EMG) transmitters affected chum salmon (Oncorhynchus keta) recovery based on three physiological parameters. Fish subjected to anesthesia plus EMG transmitter attachment (EMG group), anesthesia only (AO group), and no handling (control) were placed in a swim tunnel. Critical swimming speed (U _crit), oxygen consumption (MO₂), and muscle activity (EMG values) were assessed 0, 1, 6, 12, 24, and 30 h after treatment. The MO₂ in the EMG and AO groups was higher than in the control group 1 h after treatment, but did not differ significantly from the control in all subsequent trials (from 6 to 30 h after treatment). Values for U _crit and EMG were not significantly different from the control group in any of the trials conducted 1–30 h after treatment. We concluded that chum salmon had regained their normal swimming ability by 6 h after treatment and could be safely released into the natural environment.     

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.     

Feeding behaviour of fast- and slow-growing strains of rainbow trout, Oncorhynchus mykiss (Walbaum), during first feeding

The development of feeding behaviour in two rainbow trout strains and their hybrid was compared at the transition from endogenous nutrition to exogenous feeding. After 60 min of acclimatization, the swimming pattern, spatial distribution, agonistic behaviour and snapping of fish were quantified by making 5-min observations on groups of five fish. Behavioural patterns were recorded in four replicate groups per treatment, before and after the introduction of dry trout feed or Artemia nauplii (actometric tests) or extracts of these (olfactometric tests). Fish of the slow growing strain were less active than those of the other two strains. Greater swimming activity observed in the fish of the fast growing strain was associated with more rapid feeding and a higher number of snapping responses. Such behavioural precocity is of interest because it could provide the basis for the growth differences between the fast and slow growing strains.     

Physical activity and behavior of a centrarchid fish, Micropterus salmoides (Lacépède), during spawning

Abstract – The spawning behavior of male and female largemouth bass Micropterus salmoides (Lacépède) was studied in central Illinois during the spring of 1998 to examine patterns of muscular activity associated with different spawning related behaviors and to evaluate whether electromyogram (EMG i ) telemetry could be used to detect spawning activity. Fish were implanted with EMG i transmitters (8 females, 16 males) on April 7, prior to the initiation of spawning, and were released in four 0.10-ha earthen research ponds. Continuous EMG i records, underwater videography and additional visual observations for one pair of EMG i tagged fish were collected throughout the entire spawning event, allowing us to quantify behavioral correlates of physical activity. Male EMG i activity patterns were only correlated with female patterns during courting and periods of male aggression toward the female. Overall, EMG i activity was highest for the female during shuddering (gamete deposition), whereas male EMG i activity was similarly high during periods of nest excavation, shuddering and post-spawn parental care activities. During spawning, female EMG i activity was positively correlated to shuddering. Average daily EMG i activities for females peaked on the day of spawning. As a result of their engagement in parental care activities, male activity continued to rise even after spawning was completed. EMG i telemetry appears to be a useful technique for monitoring the reproductive activity of largemouth bass, especially in areas of high cover or turbid water or during low light conditions. This technology permitted us to quantify and contrast the activity patterns associated with different spawning related activities, information that will be required to construct accurate, gender-specific bioenergetics models for this species.     

Biochemical responses of matrinxãBrycon cephalus (Günther, 1869) after sustained swimming

Juvenile matrinxã, Brycon cephalus, were submitted to sustained swimming for 72 days at 1.0 body length s^?1. Exercised fish (EF) grew more than non‐EF and their feed conversion ratio (FCR) improved; haematological responses demonstrated a decrease in haemoglobin and mean cell haemoglobin contents and increase in the mean cell volume. In the plasma, sodium, ammonia and amino acid concentrations increased; plasma triglycerides decreased while free fatty acids increased. Liver glucose, free amino acids, ammonia, the rate protein per fish weight and total lipid content increased, while the glycogen per fish ratio declined. Glutamate dehydrogenase (GDH) activity increased while pyruvate kinase (PK) and lactate dehydrogenase (LDH) decreased. White muscle glucose, lactate, the glycogen per fish‐weight ratio and total lipid content exhibited a decrease in their values; ammonia, free amino acids and the protein per fish‐weight ratio increased. GDH and PK decreased their activities. In the red muscle glycogen store, the glycogen per fish‐weight ratio and glucose were reduced. Juvenile matrinxãs, under sustained swimming, were physiologically and biochemically adapted to exercise as indicated by improved blood flow, transport and oxygen uptake, FCR, amino acid and protein incorporation and growth. Continuous exercise is a good practice for B. cephalus cultivation.     

三疣梭子蟹配合饲料的初步研究

根据三疣梭子蟹的营养需求研制生产了硬颗粒配合饲料，并与鲜杂鱼进行养殖效果对比试验。结果表明，用颗粒饲料投喂的梭子蟹，不仅生长，蜕壳安全正常。且生长速度与鲜杂饲料投喂相当，饲料系列相当于鲜杂鱼的4-5.8倍，生产1kg梭子蟹饲料成本节约23.6%。     

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.     

Swimming behaviour as an indicator of low growth rate and impaired welfare in Atlantic halibut (Hippoglossus hippoglossus L.) reared at three stocking densities

The Atlantic halibut (Hippoglossus hippoglossus L.) is a new species in Norwegian aquaculture. However, in ongrowing trials, halibut shows variable and generally poor rates of growth. The halibut is a flatfish that spends most of its time resting on the bottom, and it was hypothesized that high stocking densities with frequent social interactions could lead to behaviour changes, reduced appetite and impaired welfare. Halibut were kept in six tanks at low, medium, and high densities (18%, 54% and 112% bottom coverage). All fish were individually tagged with Trovan™ passive implant transponder (PIT) tags, allowing an antenna to register fish swimming at the surface. Swimming and feeding behaviour was also recorded by underwater video cameras. Individual growth rates were highly variable, but food consumption and growth rates fell significantly with increasing stocking density, while individual swimming activity rose with increasing density. Frequent “surface swimmers” had a significantly lower growth rate than fish that were seldom recorded by the PIT antennae. Surface swimming may therefore be an indicator of suboptimal growth rates and impaired welfare in reared halibut.     

Fish under exercise

Improved knowledge on the swimming physiology of fish and its application to fisheries science and aquaculture (i.e., farming a fitter fish) is currently needed in the face of global environmental changes, high fishing pressures, increased aquaculture production as well as increased concern on fish well-being. Here, we review existing data on teleost fish that indicate that sustained exercise at optimal speeds enhances muscle growth and has consequences for flesh quality. Potential added benefits of sustained exercise may be delay of ovarian development and stimulation of immune status. Exercise could represent a natural, noninvasive, and economical approach to improve growth, flesh quality as well as welfare of aquacultured fish: a FitFish for a healthy consumer. All these issues are important for setting directions for policy decisions and future studies in this area. For this purpose, the FitFish workshop on the Swimming Physiology of Fish () was organized to bring together a multidisciplinary group of scientists using exercise models, industrial partners, and policy makers. Sixteen international experts from Europe, North America, and Japan were invited to present their work and view on migration of fishes in their natural environment, beneficial effects of exercise, and applications for sustainable aquaculture. Eighty-eight participants from 19 different countries contributed through a poster session and round table discussion. Eight papers from invited speakers at the workshop have been contributed to this special issue on The Swimming Physiology of Fish.     

Characterization of Gelatinolytic Activity in Seminal Plasma of Some Teleost Fish

Gelatin-containing SDS-PAGE combined with the incubation of gels in buffers containing protease inhibitors was performed for the visualization and characterization of proteolytic activity in teleost fish seminal plasma. To demonstrate the class of detected enzymes we used serine protease inhibitor – benzamidine or EDTA which inhibits metalloproteases activity. Additionally the effects of calcium ions on protease activity were investigated. Multiple gelatinolytic activities in seminal plasma of 10 teleost fish species from three orders (Cypriniformes, Salmoniformes, Perciformes) were found. Most proteases were either stimulated by Ca²⁺ and inhibited by EDTA or inhibited by benzamidine. This suggests that metalloproteases and serine proteases are major gelatinolytic proteases of fish seminal plasma. In cyprinid species we found a common profile of two gelatinolytic activities; the first band (60–66 kDa) belonged to metalloproteases, and the second one (76–81 kDa) belonged to serine proteases. Other bands were also visible and they represented mostly serine protease activity. Species from the Salmoniformes order showed a similarity in metalloproteases with molecular weights of about 64 and 75 kDa. Salmonid species also had similar serine proteases with molecular weights of about 102 and 165 kDa. In European grayling seminal plasma we found metalloproteases with molecular weight of 51, 57, 64, 70 kDa and two serine proteases activities of 35 and 125 kDa. Percid species had metalloproteases activities of 53 and 63 kDa and serine protease activity of 100 kDa. Protease of other, presently unknown classes were also found in seminal plasma of asp, chub, European grayling and pikeperch. The physiological role of seminal plasma proteases is still unknown.     

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.