David J. Ellerby

Sympatric Divergence and Performance Trade-Offs of Bluegill Ecomorphs

Phenotypic plasticity in response to environmental cues can create distinct morphological types within populations. This variation in form, and potentially function, may be a factor in initiating population divergence and the formation of new species. Here we show the translation of sympatric, habitat-specific morphological divergence into performance differences in energy economy, maneuverability and steady-state locomotion. Littoral and pelagic bluegill sunfish ecomorphs show differences in performance that appear adaptive within their respective habitats: greater maneuverability in the heavily vegetated littoral; greater steady-state swimming speed and economy in the open-water pelagic. This represents a trade-off in unsteady versus steady swimming performance, likely because morphological features associated with maximizing maneuverability are incompatible with enhancing steady-swimming performance. This may constrain the direction of adaptive change, maintaining the divergence created by phenotypic plasticity. The combination of habitat specific sympatric adaptation and constraints imposed by performance trade-offs may be an important factor underlying the high rate of speciation in freshwater fishes from post-glacial lakes.

The mechanics of explosive seed dispersal in orange jewelweed (Impatiens capensis)

Explosive dehiscence ballistically disperses seeds in a number of plant species. During dehiscence, mechanical energy stored in specialized tissues is transferred to the seeds to increase their kinetic and potential energies. The resulting seed dispersal patterns have been investigated in some ballistic dispersers, but the mechanical performance of a launch mechanism of this type has not been measured. The properties of the energy storage tissue and the energy transfer efficiency of the launch mechanism were quantified in Impatiens capensis. In this species the valves forming the seed pod wall store mechanical energy. Their mass specific energy storage capacity (124u2009J kg−1) was comparable with that of elastin and spring steel. The energy storage capacity of the pod tissues was determined by their level of hydration, suggesting a role for turgor pressure in the energy storage mechanism. During dehiscence the valves coiled inwards, collapsing the pod and ejecting the seeds. Dehiscence took 4.2±0.4u2009ms (mean ±SEM, n=13). The estimated efficiency with which energy was transferred to the seeds was low (0.51±0.26%, mean ±SEM, n=13). The mean seed launch angle (17.4±5.2, mean ±SEM, n=45) fell within the range predicted by a ballistic model to maximize dispersal distance. Low ballistic dispersal efficiency or effectiveness may be characteristic of species that also utilize secondary seed dispersal mechanisms.

Swimming Flumes as a Tool for Studying Swimming Behavior and Physiology: Current Applications and Future Developments

Swimming flumes enable fish swimming behavior, physiology, and performance to be quantified in ways that are not practicable for fish swimming through open water. By placing fish in a water flow, speed can be controlled, fish can be instrumented to monitor a wide range of physiological parameters, and the exchange of materials between the fish and water can be quantified. This can provide vital information regarding fish fitness and health. If meaningful data are to be obtained, however, careful consideration must be given to flume design and operation, experimental protocol and the physiological state of the fish. Modifications to standard flume designs can potentially allow for accommodation of a wider range of species and experimental conditions that will enhance basic understanding of fish physiology and behavior and can potentially be applied in optimizing aquacultural techniques.

The seed dispersal catapult of Cardamine parviflora (Brassicaceae) is efficient but unreliable

UNLABELLEDnnnnPREMISE OF THE STUDYnSeed dispersal performance is an essential component of plant fitness. Despite their significance in shaping performance, the mechanical processes that drive dispersal are poorly understood. We have quantified seed dispersal mechanics in Cardamine parviflora (Brassicaceae), a ballistic disperser that launches seeds with specialized catapult-like structures. To determine which aspects of catapult function dictate interspecific dispersal differences, we compared this disperser with other ballistic dispersers. Comparison with brassicas that lack ballistic dispersal may also provide insight into the evolution of this mechanism. •nnnMETHODSnCatapult performance was quantified using high-speed video analysis of dehiscence, ballistic modeling of seed trajectories, and measuring the mechanical energy storage capacity of the spring-like siliqua valve tissue that launched the seeds. •nnnKEY RESULTSnThe siliquae valves coiled rapidly outward, launching the seeds in 4.7 ± 1.3 ms (mean ± SD, N = 11). Coiling was likely driven by the bilayered valve structure. The catapult was 21.3 ± 10.3% efficient (mean ± SD, N = 11) at transferring stored elastic energy to the seeds as kinetic energy. The majority of seeds (71.4%) were not launched effectively. •nnnCONCLUSIONSnThe efficiency of the C. parviflora catapult was high in comparison to that of a ballistic diplochore, a dispersal mode associated with poor ballistic performance, although the unreliability of the launch mechanism limited dispersal distance. Effective launching requires temporary seed-valve adhesion. The adhesion mechanism may be the source of the unreliability. Valve curvature is likely driven by the bilayered valve structure, a feature absent in nonballistic brassicas.

Serotonin as an integrator of leech behavior and muscle mechanical performance.

The obliquely striated muscle in the leech body wall has a broad functional repertoire; it provides power for both locomotion and suction feeding. It also operates over an unusually high strain range, undergoing up to threefold changes in length. Serotonin (5-HT) may support this functional flexibility, integrating behavior and biomechanics. It can act centrally, promoting motor outputs that drive body wall movements, and peripherally, modulating the mechanical properties of body wall muscle. During isometric contractions 5-HT enhances active force production and reduces resting muscle tone. We therefore hypothesized that 5-HT would increase net work output during the cyclical contractions associated with locomotion and feeding. Longitudinal strains measured during swimming, crawling and feeding were applied to body wall muscle in vitro with the timing and duration of stimulation selected to maximize net work output. The net work output during all simulated behaviors significantly increased in the presence of 100μM 5-HT relative to the 5-HT-free control condition. Without 5-HT the muscle strips could not achieve a net positive work output during simulated swimming. The decrease in passive tension associated with 5-HT may also be important in reducing muscle antagonist work during longitudinal muscle lengthening. The behavioral and mechanical effects of 5-HT during locomotion are clearly complementary, promoting particular behaviors and enhancing muscle performance during those behaviors. Although 5-HT can enhance muscle mechanical performance during simulated feeding, low in vivo activity in serotonergic neurons during feeding may mean that its mechanical role during this behavior is less important than during locomotion.

Trade-offs between performance and variability in the escape responses of bluegill sunfish (Lepomis macrochirus)

Successful predator evasion is essential to the fitness of many animals. Variation in escape behaviour may be adaptive as it reduces predictability, enhancing escape success. High escape velocities and accelerations also increase escape success, but biomechanical factors likely constrain the behavioural range over which performance can be maximized. There may therefore be a trade-off between variation and performance during escape responses. We have used bluegill sunfish (Lepomis macrochirus) escape responses to examine this potential trade-off, determining the full repertoire of escape behaviour for individual bluegill sunfish and linking this to performance as indicated by escape velocity and acceleration. Fish escapes involve an initial C-bend of the body axis, followed by variable steering movements. These generate thrust and establish the escape direction. Directional changes during the initial C-bend were less variable than the final escape angle, and the most frequent directions were associated with high escape velocity. Significant inter-individual differences in escape angles magnified the overall variation, maintaining unpredictability from a predator perspective. Steering in the latter stages of the escape to establish the final escape trajectory also affected performance, with turns away from the stimulus associated with reduced velocity. This suggests that modulation of escape behaviour by steering may also have an associated performance cost. This has important implications for understanding the scope and control of intra- and inter-individual variation in escape behaviour and the associated costs and benefits.

The effects of steady swimming on fish escape performance.

Escape maneuvers are essential to the survival and fitness of many animals. Escapes are frequently initiated when an animal is already in motion. This may introduce constraints that alter the escape performance. In fish, escape maneuvers and steady, body caudal fin (BCF) swimming are driven by distinct patterns of curvature of the body axis. Pre-existing muscle activity may therefore delay or diminish a response. To quantify the performance consequences of escaping in flow, escape behavior was examined in bluegill sunfish (Lepomis macrochirus) in both still-water and during steady swimming. Escapes executed during swimming were kinematically less variable than those made in still-water. Swimming escapes also had increased response latencies and lower peak velocities and accelerations than those made in still-water. Performance was also lower for escapes made up rather than down-stream, and a preference for down-stream escapes may be associated with maximizing performance. The constraints imposed by pre-existing motion and flow, therefore, have the potential to shape predator–prey interactions under field conditions by shifting the optimal strategies for both predators and prey.

Field swimming behavior in largemouth bass deviates from predictions based on economy and propulsive efficiency

ABSTRACT Locomotion is energetically expensive. This may create selection pressures that favor economical locomotor strategies, such as the adoption of low-cost speeds and efficient propulsive movements. For swimming fish, the energy expended to travel a unit distance, or cost of transport (COT), has a U-shaped relationship to speed. The relationship between propulsive kinematics and speed, summarized by the Strouhal number (St=fA/U, where f is tail beat frequency, A is tail tip amplitude in m and U is swimming speed in m s−1), allows for maximal propulsive efficiency where 0.2<St<0.4. Largemouth bass adopted field speeds that were generally below the range predicted to minimize their COT. This may reflect speed modulation to meet competing functional demands such as enabling effective prey detection and capture. St exceeded the optimal range for the lowest observed swimming speeds. Mechanical and physiological constraints may prevent adoption of efficient St during low-speed swimming. Highlighted Article: Video analysis of largemouth bass swimming behavior in the field suggests that they do not use swimming speeds or propulsive movements that maximize economy and efficiency.

Field swimming performance of bluegills sunfish, Lepomis macrochirus: implications for field activity cost estimates and laboratory measures of swimming performance

Abstract Mobility is essential to the fitness of many animals, and the costs of locomotion can dominate daily energy budgets. Locomotor costs are determined by the physiological demands of sustaining mechanical performance, yet performance is poorly understood for most animals in the field, particularly aquatic organisms. We have used 3‐D underwater videography to quantify the swimming trajectories and propulsive modes of bluegills sunfish (Lepomis macrochirus, Rafinesque) in the field with high spatial (1–3 mm per pixel) and temporal (60 Hz frame rate) resolution. Although field swimming trajectories were variable and nonlinear in comparison to quasi steady‐state swimming in recirculating flumes, they were much less unsteady than the volitional swimming behaviors that underlie existing predictive models of field swimming cost. Performance analyses suggested that speed and path curvature data could be used to derive reasonable estimates of locomotor cost that fit within measured capacities for sustainable activity. The distinct differences between field swimming behavior and performance measures obtained under steady‐state laboratory conditions suggest that field observations are essential for informing approaches to quantifying locomotor performance in the laboratory.

Linking muscle metabolism and functional variation to field swimming performance in bluegill sunfish (Lepomis macrochirus)

Skeletal muscle has diverse mechanical roles during locomotion. In swimming fish, power-producing muscles work in concert with the accessory muscles of the fins which augment and control power transfer to the water. Although fin muscles represent a significant proportion of the locomotor muscle mass, their physiological properties are poorly characterized. To examine the relationship between muscle metabolism and the differing mechanical demands placed on distinct muscle groups, we quantified the aerobic and glycolytic capacities of the myotomal, pectoral and caudal muscles of bluegill sunfish. These were indicated by the activities of citrate synthase and lactate dehydrogenase, rate-limiting enzymes for aerobic respiration and glycolysis, respectively. The well-established roles of slow and fast myotomal muscle types in sustained and transient propulsive movements allows their use as benchmarks to which other muscles can be compared to assess their function. Slow myotomal muscle had the highest CS activity, consistent with meeting the high metabolic and mechanical power demands of body-caudal fin (BCF) swimming at the upper end of the aerobically supported speed range. The largest pectoral adductors and abductors had CS activities lower than the slow myotomal muscle, in line with their role supplying thrust for low-speed, low-power swimming. The metabolic capacities of the caudal muscles were surprisingly low and inconsistent with their activity during steady-state BCF swimming at high speeds. This may reflect adaptation to the observed swimming behavior in the field, which typically involved short bouts of BCF-propulsive cycles rather than sustained propulsive activity.