Morphological correlates of swimming activity in wild largemouth bass (Micropterus salmoides) in their natural environment.

Individual variation in morphology has been linked to organismal performance in numerous taxa. Recently, the relationship between functional morphology and swimming performance in teleost fishes has been studied in laboratory experiments. In this study, we evaluate the relationship between morphology and swimming activity of wild largemouth bass (Micropterus salmoides) during the reproductive period, providing the first data derived on free-swimming fish not exposed to forced swim trials in the laboratory. Sixteen male largemouth bass were angled from their nests, telemetered, and subsequently monitored by a whole-lake acoustic hydrophone array with sub-meter accuracy. Additionally, eleven morphological measurements were taken from digital images of each fish. A principal components analysis of the morphological measurements described 79.8% of the variance. PC1 was characterized by measures of overall body stoutness, PC2 was characterized by measures of the length and depth of the caudal region, and PC3 characterized individuals with relatively large anterior portions of the body and relatively small caudal areas. Of these variables, only PC3 showed significant relationships to swimming activity throughout the parental care period. PC3 was negatively correlated with multiple measures of swimming activity across the parental care period. Furthermore, swimming performance of individual male bass was noted to be repeatable across the parental care period indicating that this phenomenon extends beyond the laboratory.     

Morphological correlates of swimming activity in wild largemouth bass (Micropterus salmoides) in their natural environment

Individual variation in morphology has been linked to organismal performance in numerous taxa. Recently, the relationship between functional morphology and swimming performance in teleost fishes has been studied in laboratory experiments. In this study, we evaluate the relationship between morphology and swimming activity of wild largemouth bass (Micropterus salmoides) during the reproductive period, providing the first data derived on free-swimming fish not exposed to forced swim trials in the laboratory. Sixteen male largemouth bass were angled from their nests, telemetered, and subsequently monitored by a whole-lake acoustic hydrophone array with sub-meter accuracy. Additionally, eleven morphological measurements were taken from digital images of each fish. A principal components analysis of the morphological measurements described 79.8% of the variance. PC1 was characterized by measures of overall body stoutness, PC2 was characterized by measures of the length and depth of the caudal region, and PC3 characterized individuals with relatively large anterior portions of the body and relatively small caudal areas. Of these variables, only PC3 showed significant relationships to swimming activity throughout the parental care period. PC3 was negatively correlated with multiple measures of swimming activity across the parental care period. Furthermore, swimming performance of individual male bass was noted to be repeatable across the parental care period indicating that this phenomenon extends beyond the laboratory.     

A reappraisal of activity metabolism in Atlantic cod (Gadus morhua)

Atlantic cod ( Gadus morhua ) were forced to swim in a swim tunnel respirometer until fatigued; oxygen consumption rate (O₂) was measured during swimming at incremental speeds until the fish was exhausted and during recovery from exhaustion. Maximal oxygen consumption (O_2max) occurred during maximal activity as has been found for other fish species, but at odds with the current paradigm for Atlantic cod. Earlier experiments had drawn the conclusion that O_2max in Atlantic cod occurs during recovery from exhaustive exercise. We found no support for this paradigm in our experiments and we propose that the respiratory physiology of Atlantic cod is not unlike that of other fishes.     

Empirical analysis of the influence of swimming pattern on the net energetic cost of swimming in fishes

We tested the hypothesis that the energetics of swimming in a flume accurately represent the costs of various spontaneous movements using empirical relationships between fish swimming costs, weight, and speed for three swimming patterns: (1) 'forced swimming' corresponded to movements adopted by fish forced to swim against a unidirectional current of constant velocity; (2) 'directed swimming' was defined as quasi-rectilinear movements executed at relatively constant speeds in a stationary body of water and (3) 'routine swimming' was characterized by marked changes in swimming direction and speed. Weight and speed explained between 76% (routine swimming) and 80% (forced swimming) of net swimming cost variability. Net costs associated with different swimming patterns were compared using ratios of model predictions (swimming cost ratio; SCR) for various weight and speed combinations. Routine swimming was the most expensive swimming pattern (SCR for routine and forced swimming =6.4 to 14.0) followed by directed (SCR for directed and forced swimming =0.9 to 2.8), and forced swimming. The magnitude of the difference between the net costs of forced and spontaneous swimming increases with movement complexity and decreases as fish weight increases.     

Activity patterns of juvenile Atlantic cod (Gadus morhua) in Buckley Cove,Newfoundland

We monitored swimming speed of 2–3 year-old juvenile Atlantic cod (Gadus morhua) from August to December 1999, using a 2-D location finding acoustic telemetry system in a coastal area of Newfoundland, Canada. We concurrently monitored the locations of 22–41 individuals by triangulation using a fixed hydrophone array. We estimated average swimming speeds at intervals of 60–120 s and compared them over a 1 to 17 °C thermal range, three diel periods, and five substrates (sand, gravel, sand-sparse boulder, boulder, and kelp). However, cod did not exhibit a change in swimming speed over the temperature range studied. Increased activity and foraging rates (expressed as swimming speeds) were expected to increase at elevated temperatures due to increased metabolic demands. Activity did vary significantly with diel cycle and substrate. Swimming speeds were significantly lower at night during September and October. Results for August and November were inconclusive, while swimming speed was significantly lower during the day in December. We observed significantly reduced average swimming speeds in structurally complex substrates (e.g. rock, cobble and kelp) in September and October. Our results suggest that activity of juvenile cod in the wild does not vary with temperature as predicted from studies in the laboratory. Instead, activity varied with diel cycles and structural complexity, variables that influence an individual's ability to forage and seek refuge, potentially altering individual fitness.     

Diel activity pattern and food search behaviour in cod, Gadus morhua

Food search behaviour in cod, Gadus morhua, was studied by means of a stationary positioning system. Six fish in a fjord in northern Norway were tagged in situ by allowing them to shallow bait-wrapped acoustic tags, and their swimming behaviour was continuously recorded. The fish exhibited a diel activity rhythm, with higher swimming speeds and a larger range during the day than at night. Chemically mediated food search was studied in periods of both high and low activity by setting a string of baits in the morning and at night. During the period of high activity, more fish localized the baits (70 versus 45% of the observations) and the time to localization was 50% shorter, indicating that high swimming activity increased the probability of encountering the odour plume and the odour source. No diel variations in the response threshold to olfactory stimuli were found, as the proportion of fish in the odour plume that responded rheotactically, and their response distances were similar in the two periods (240 and 227 m respectively). Cod performed area-restricted searches on encountering the string of baits during the day. The probability of cod detecting prey by taste receptors, encountering the odour plume of prey or localizing a stationary food source after olfactory stimulation should be relatively independent of light, and this could explain why cod were active throughout the 24 h cycle. On the other hand, the visual detection distance and the ability to capture active prey are influenced by the light level, resulting in higher activity during the day.     

Oxygen consumption of active rainbow trout, Salmo gairdneri Richardson, derived from electromyograms obtained by radiotelemetry

A radiotelemetry apparatus is described for sensing and transmitting electromyograms (EMGs) from free-swimming fish. EMGs are recorded from the epaxial muscles of adult rainbow trout during periods in spontaneous (= routine) activity, and forced-swim, respirometers. When such EMG records are integrated, subjected to spectral analysis, and computer-averaged, the EMG values (in μV) are highly correlated with the fish oxygen consumption during the activity periods. However, there is a marked difference between the regression slopes for oxygen v . EMG value for the data from the spontaneous, and forced-swim, respirometers; the former slope is the steeper. The probable explanation of this phenomenon is that whereas in forced swims the epaxial myomeres are responsible for most of the activity of the fish, in spontaneous activity other muscle systems (e.g. of the lateral, dorsal and ventral fins) come to account for a greater relative proportion of body movement. The difference in slope, although great, is evidently a regular phenomenon. The shift from one regression to the other occurs at a fairly precise epaxial EMG value ( c . 5 μV). This suggests that the laboratory calibration of EMG value in terms of oxygen consumption can be utilized in the wild so that EMG records from free-swimming fish, fitted with telemetry packages can be used to deduce oxygen consumption attributable to activity. It also appears that such records can be used as a guide to the type of activity of the fish, i.e. desultory movements or free cruising.     

Effects on temperature on spontaneous and thyroxine-stimulated locomotor activity of Atlantic cod

Spontaneous locomotor activity of cod Gadus morhua maintained at 6° C tripled from February to May. In contrast, locomotor activity of cod held at 2° C was significantly lower than at 6° C (between 25 and 65% lower) and the seasonal increase was smaller. Plasma levels of both thyroxine (T₄) and triiodothyronine (T₃) did not differ between 2 and 6° C. T₄ injection increased locomotor activity by 10% for both temperature regimes. These data indicate that low water temperature reduces locomotor activity associated with migration in cod and that thyroid hormones are not involved in this decrease. This study provides a possible mechanism through which cold waters may affects migration and distribution of cod via its Effects on locomotor activity and swimming speed.     

Effects of turbidity on the spontaneous and prey-searching activity of juvenile Atlantic cod (Gadus morhua)

Increasing turbidity in coastal waters in the North Atlantic and adjacent seas has raised concerns about impacts on Atlantic cod (Gadus morhua) using these areas as nurseries. A previous experiment (Meager et al. 2005 Can. J. Fish. Aquat. Sci. 62, 1978-1984) has shown that turbidity (up to 28 beam attenuation m-1) had little effect on the foraging rate of juvenile cod. Although this was attributed to cod using chemoreception in conjunction with vision to locate prey, foraging rates may also be maintained by increased activity. Higher activity, however, is energetically costly and may offset benefits from increased foraging return. We examined the effects of turbidity on prey searching and spontaneous activity of juvenile cod in the laboratory, by measuring activity with and without prey cues. Activity of juvenile cod was nonlinearly affected by turbidity and was lower at intermediate turbidity, regardless of the presence of prey odour. Activity increased over time when prey odour was present and decreased when absent, but the effects of prey odour were similar across all turbidity levels. Position in the tank was unaffected by turbidity or prey odour. Reduced activity at intermediate turbidities is likely to offset longer prey-search times. At high turbidity (greater than 17m-1), both longer prey-search times and higher activity indicate that increased energetic costs are probable.     

Opercular differential pressure as a predictor of metabolic oxygen demand in the starry flounder

The feasibility of using a differential pressure sensor connected to an acoustic telemetry device to monitor opercular activity as a correlate of oxygen consumption was investigated. Four starry flounders Platichthys stellatus were fitted with a miniature differential pressure sensor mounted close to the operculum. A cannula was connected to the sensor and inserted under the operculum, inside the branchial cavity. Measurements of oxygen consumption and opercular activity were carried out over a broad range of metabolic activity, from the post‐surgery stress (high metabolic rate) to routine metabolic rate the following day. Relationships between differential pressure changes (rate and amplitude) were highly correlated with oxygen consumption ( r ² = 0·74 and 0·60 respectively). The results indicate that monitoring opercular activity offers an alternative method for measuring aerobic metabolism in free‐swimming fishes in nature.     

Nitric oxide is not involved in the control of vasopressin release during acute forced swimming in rats

Summary. Neurons of the hypothalamo-neurohypophyseal system (HNS) are known to contain high amounts of neuronal nitric oxide (NO) synthase (nNOS). NO produced by those neurons is commonly supposed to be involved as modulator in the release of the two nonapeptides vasopressin (AVP) and oxytocin into the blood stream. Previous studies showed that forced swimming fails to increase the release of AVP into the blood stream while its secretion into the hypothalamus is triggered. We investigated here whether hypothalamically acting NO contributes to the control of the AVP release into blood under forced swimming conditions. Intracerebral microdialysis and in situ hybridization were employed to analyze the activity of the nitrergic system within the supraoptic nucleus (SON), the hypothalamic origin of the HNS. A 10-min forced swimming session failed to significantly alter the local NO release as indicated both by nitrite and, the main by-product of NO synthesis, citrulline levels in microdialysis samples collected from the SON. Microdialysis administration of NO directly into the SON increased the concentration of AVP in plasma samples collected during simultaneous forced swimming. In an additional experiment the effect of the defined stressor exposure on the concentration of mRNA coding for nNOS within the SON was investigated by in situ hybridization. Forced swimming increased the expression of nNOS mRNA at two and four hours after onset of the stressor compared to untreated controls. Taken together, our results imply that NO within the SON does not contribute to the regulation of the secretory activity of HNS neurons during acute forced swimming. Increased nNOS mRNA in the SON after forced swimming and the increase in AVP release in the presence of exogenous NO under forced swimming points to a possible role of NO in the regulation of the HNS under repeated stressor exposure.Current address: Departments of Behavioral Neuroscience and Neurology, Oregon Health & Science University, Portland, OR 97239, U.S.A.     

A biotelemetry system recording fish activity

A biotelemetry system is described for obtaining, transmitting and recording the electromyograms (EMGs) produced in muscle activity of free-swimming fish as quantitative indicators of overall fish activity. The radiotransmitters used come in the form of cylindrical packages having two sensing electrodes, all fully implantable in the fish body cavity. EMGs are transmitted as radio pulses with the intensity of muscular activity determining the intervals between pulses. The packages also contain temperature sensors and fish temperatures are transmitted with every 32nd pulse. Transmitted EMG pulses are detected, 'measured' and stored by a single portable receiver (Model SRX_400, Lotek). Data can be subsequently transferred to a computer (which can also be portable) for storage, processing and statistical analysis. Transmitter battery life can be in excess of 7 months, permitting laboratory or field studies of long duration. Transmitter package implantation surgery requires a mid-ventral incision and internal securing of transmitter and sensing electrodes. Surgical silk, cyanoacrylate tissue adhesives, and polydioxanone (PD), a synthetic absorbable suture, were all tried as means of incision closure. The most effective was PD alone. Trials of the system consisted of forced swims by transmitter-equipped rainbow trout Oncorhynchus mykiss Walbaum. The data obtained provided an inverse linear relation between forced swim speed and EMG pulse interval. Trials were conducted at intervals over periods up to 2 months. Fish showed neither distress, nor difficulty in swimming up to maximum speeds of 60 cm s ⁻¹ (fish lengths 41.0, 44.4 cm).     

Estimating the active metabolic rate (AMR) in fish based on tail beat frequency (TBF) and body mass

Tail beat frequency (TBF) was measured for carp (Cyprinus carpio) and roach (Rutilus rutilus), during steady swimming at five different speeds and for fish of various body masses. A multiple stepwise linear regression analysis resulted in models for the prediction of TBFs depending on swimming speed as an independent variable. Speed explained 72 and 86% of the variance in TBF for carp and roach, respectively. By using these data to predict TBF from speed and substituting values into a model from a previous study that predicts active metabolic rates (AMR) from body mass and swimming speed, we can calculate AMR from only fish mass and TBF. Thus, the derived models can be used to estimate the AMR in fish by measuring TBFs in the field using biotelemetry. The approach presented here is a useful and relatively simple tool for estimating the activity metabolism in free-swimming fish. In future studies this method should be applied to a larger and more representative sample size to test the applicability and the validity for a broader range of species.     

Power isn't everything: muscle function and energetic costs during steady swimming in Atlantic cod (Gadus morhua)

Power produced by red myotomal muscles of fish during cruise swimming appears seldom maximized, so we sought to investigate whether economy may impact or dominate muscle function. We measured cost of transport (COT) using oxygen consumption and the strain trajectories and electromyographic activity of red muscle measured at anterior (ANT) and posterior (POST) locations while Atlantic cod (Gadus morhua) swam steadily at speeds between 0.3 and 1.0 body lengths (BL) s(-1). We then measured the power produced by isolated segments of red muscle when activated either as in the swimming cod or such that maximal net power was produced. Patterns of activation during swimming were not optimal for power output and were highly variable between tail beats, particularly at the ANT location and at slow swim speeds. Muscle strain amplitude did not increase until swimming speed reached 0.9 (ANT) versus 0.5 (POST) BL s(-1). These limited power to only 53% (ANT) and 71% (POST) of maximum at slower swim speeds and to 70%-80% of maximum at high swim speeds. COT (resting metabolism subtracted) was minimal at the slowest swim speed, surprisingly, where power was most impaired by activation and strain. Thus, production of powered forces for maneuverability/stability appeared to greatly impact red muscle function during cruise swimming in cod, particularly at slow speeds and in ANT muscle.     

Effects of endogenous rhythms and light conditions on foraging and predator-avoidance in juvenile plaice

The influences of a light: dark cycle and a persistent endogenous rhythm of activity on foraging (on the bivalve Donax vittatus ) and avoiding a predator (juvenile cod, Gadus morhua ) were investigated in freshly-caught juvenile plaice, Pleuronectes plalessa . Time lapse video recordings were made of fish in the presence and absence of prey and predators in laboratory tanks over 24-hour periods between the times of successive daytime low waters. Endogenous rhythms of activity were seen in all experimental treatments. Swimming both close to the bottom and in the water column showed a strong circatidal rhythm, with most activity 2 to 3 h after the predicted time of high water. Swimming in the water column was more frequent at night than by day. In the presence of a population of Donax , whose siphon tips could be eaten as food, swimming close to the bottom became more frequent. This increase in benthic swimming was independent of the endogenous cycle of activity and was correlated with the frequency of attacks on siphons. The presence of the cod predator delayed the onset of foraging activity, producing a foraging/predator avoidance trade-off. The independence of foraging from light and endogenous rhythms suggests that this trade-off may be similarly independent. The cod also greatly reduced swimming in the water column in darkness, behaviour apparently unrelated to foraging.     

Interaction between hindbrain and spinal networks during the development of locomotion in zebrafish

Little is known about the role of the hindbrain during development of spinal network activity. We set out to identify the activity patterns of reticulospinal (RS) neurons of the hindbrain in fictively swimming (paralyzed) zebrafish larvae. Simultaneous recordings of RS neurons and spinal motoneurons revealed that these were coactive during spontaneous fictive swim episodes. We characterized four types of RS activity patterns during fictive swimming: (i) a spontaneous pattern of discharges resembling evoked high-frequency spiking during startle responses to touch stimuli, (ii) a rhythmic pattern of excitatory postsynaptic potentials (EPSPs) whose frequency was similar to the motoneuron EPSP frequency during swim episodes, (iii) an arrhythmic pattern consisting of tonic firing throughout swim episodes, and (iv) RS cell activity uncorrelated with motoneuron activity. Despite lesions to the rostral spinal cord that prevented ascending spinal axons from entering the hindbrain (normally starting at approximately 20 h), RS neurons continued to display the aforementioned activity patterns at day 3. However, removal of the caudal portion of the hindbrain prior to the descent of RS axons left the spinal cord network unable to generate the rhythmic oscillations normally elicited by application of N-methyl-d-aspartate (NMDA), but in approximately 40% of cases chronic incubation in NMDA maintained rhythmic activity. We conclude that there is an autonomous embryonic hindbrain network that is necessary for proper development of the spinal central pattern generator, and that the hindbrain network can partially develop independently of ascending input.     

Field metabolic rates of rainbow trout estimated using electromyogram telemetry

Field metabolic rates (FMR) of five rainbow trout were estimated using electromyogram (EMG) telemetry of the axial muscle. A series of laboratory experiments indicated that the EMG transmitter output was related strongly to total oxygen consumption of the fish over a wide range of swimming speeds and temperatures. No differences were evident when the oxygen consumption v . EMG relationship for routine swimming was compared with that for forced swimming. FMR was assessed on two time scales, revealing diel patterns and seasonal patterns. On the diel scale, the FMR pattern could be classified as crepuscular. At the upper and lower limits of temperature tolerance, the diel pattern was less distinct. On the seasonal scale, mean daily FMR was strongly dependent on mean environmental temperature. Comparisons between FMR and laboratory derived estimates of standard and maximally active metabolism indicate that the rainbow trout in the field utilize <20% of the available scope for activity.     

Burst and coast use, swimming performance and metabolism of Atlantic cod Gadus morhua in sub-lethal hypoxic conditions

Prolonged swimming capacity (critical swimming speed, U _crit, protocol) and metabolism were measured for 14 Atlantic cod Gadus morhua exposed to seven oxygen levels within the non-lethal range normally encountered in the Gulf of St Lawrence (35 to 100% saturation). Burst-and-coast swimming was triggered earlier (at lower speeds) in hypoxia, and burst-and-coast movements were more frequent in hypoxia than in normoxia at low speeds. Furthermore, the metabolic scope beyond the metabolic rate at which Atlantic cod resorted to burst-and-coast movements decreased gradually as ambient oxygen concentration dropped. Overall, fewer burst-and-coast movements were observed in hypoxia while the distance swum in burst-and-coast mode remained c . 1% of the total distance swam in all tests. Oxygen availability had no effect on the rate of increase in metabolic rate with increasing velocity <50 cm s⁻¹_, but limited swimming performances and metabolic rate at higher speeds. The prevailing low oxygen tensions on the bottom in the deep channels may impair the swimming capacity of Atlantic cod in the estuary and northern Gulf of St Lawrence.     

Flagellar waveforms of gametes in the brown alga Ectocarpus siliculosus

Brown algae are members of the Stramenopiles and their gametes generally have two heterogeneous flagella: a long anterior flagellum (AF) with mastigonemes and a short posterior flagellum (PF). In this study, swimming paths and flagellar waveforms in free-swimming and thigmotactic-swimming male and female gametes and in male gametes during chemotaxis, were quantitatively analysed in the model brown alga Ectocarpus siliculosus. This analysis was performed using a high-speed video camera. It was revealed that the AF plays a role in changing the locomotion of male and female gametes from free-swimming to thigmotactic-swimming and also in changing the swimming path of male gametes from linear to circular during chemotaxis. In the presence of a sex pheromone, male gametes changed their swimming path from linear (swimming path curvature, 0–0.02 µm^–1) to middle and small circular path (swimming path curvature, 0.04–0.20 µm^–1). The flagellar asymmetry and the deflection angle of the AF became larger, whereas the oscillation pattern of the AF was stable. However, there was no correlation between the flagellar asymmetry and the deflection angle of the AF and the path curvature when the male gametes showed middle to small circular paths. The PF irregularly changed the deflection angle and the oscillation pattern was unstable depending on the gradient of the sex pheromone concentration. AF waveforms were independent of PF locomotion during chemotaxis. This means that the AF has the ability to change the swimming path of male gametes – for example, from a highly linear path to a circular path – while changes in locomotion from a middle circle path to a small circle path is the result of beating of the PF.     

Behaviour of rainbow trout Oncorhynchus mykiss presented with a choice of normoxia and stepwise progressive hypoxia

The objective of this study was to identify behavioural adjustments leading to avoidance of hypoxia. Using the oxygen-sensitive species rainbow trout Oncorhynchus mykiss as a model, individual fish were recorded while moving freely between two sides of a test arena: one with normoxia and one with stepwise progressive hypoxia [80-30% dissolved oxygen (DO) air saturation]. The results demonstrated a gradual decrease in the total time spent in hypoxia starting at 80% DO air saturation. At this DO level, the avoidance of hypoxia could not be attributed to changes in spontaneous swimming speed, neither in normoxia nor in hypoxia. Reducing the DO level to 60% air saturation resulted in decreased spontaneous swimming speed in normoxia, yet the number of trips to the hypoxic side of the test arena remained unchanged. Moreover, data revealed increased average residence time per trip in normoxia at DO levels ≤60% air saturation and decreased average residence time per trip in hypoxia at DO levels ≤50% air saturation. Finally, the spontaneous swimming speed in hypoxia increased at DO levels ≤40% air saturation and the number of trips to hypoxia decreased at the 30% DO air saturation level. Thus, avoidance of the deepest hypoxia was connected with a reduced number of trips to hypoxia as well as decreased and increased spontaneous swimming speed in normoxia and hypoxia, respectively. Collectively, the data support the conclusions that the mechanistic basis for avoidance of hypoxia may (1) not involve changes in swimming speed during mild hypoxia and (2) depend on the severity of hypoxia.