The flow field around a freely swimming copepod in steady motion. Part II: Numerical simulation

Three-dimensional, numerical simulations of the flow field arounda freely swimming model-copepod were performed using a finite-volumecode. The model copepod had a realistic body shape representedby a curvilinear body-fitted coordinate system. The beatingmovement of the cephalic appendages was replaced by a distributedforce field acting on the water ventrally adjacent to the copepod'sbody. In the simulations, we took into account that freely swimmingcopepods are self-propelled bodies through properly couplingthe Navier–Stokes equations with the dynamic equationfor the copepod's body. Flow fields were calculated for fivesteady motions: (1) hovering, (2) sinking, (3) upwards swimming,(4) backwards swimming and (5) forwards swimming. The numericalresults confirm the conclusions drawn from the theoretical analysisusing Stokes flow models by Jiang et al. [in a companion paper(Jiang et al., 2002a)] for a spherical copepod shape and showthat the geometry of the flow field around a freely swimmingcopepod varies significantly with the different swimming behaviours.When a copepod hovers in the water, or swims very slowly, itgenerates a cone-shaped and wide flow field. In contrast, whena copepod sinks, or swims fast, the flow geometry is not cone-shaped,but cylindrical, narrow and long. The relationships betweencopepods' swimming behaviour and body orientation, hydrodynamicconspicuousness, energetics as well as feeding efficiency werediscussed, based on the simulation data. It is shown that thebehaviour of hovering or swimming slowly is more energeticallyefficient in terms of relative capture volume per energy expendedthan the behaviour of swimming fast, i.e. for a same amountof energy expended a hovering or slow-swimming copepod is ableto scan more water than a fast-swimming one. The numerical resultsalso suggest that the flow field generated by a fast-swimmingcopepod enables the copepod to use mechanoreception to perceivethe food/prey and therefore increases the food concentrationin the swept volume and that the flow field around a free-sinkingcopepod favours the copepod's mechanoreception while minimizingthe energy expense, so that the energy budget can still be maintainedfor both cases.     

Copepod flow modes and modulation: a modelling study of the water currents produced by an unsteadily swimming copepod

Video observation has shown that feeding-current-producing calanoid copepods modulate their feeding currents by displaying a sequence of different swimming behaviours during a time period of up to tens of seconds. In order to understand the feeding-current modulation process, we numerically modelled the steady feeding currents for different modes of observed copepod motion behaviours (i.e. free sinking, partial sinking, hovering, vertical swimming upward and horizontal swimming backward or forward). Based on observational data, we also reproduced numerically a modulated feeding current associated with an unsteadily swimming copepod. We found that: (i) by changing its propulsive force, a copepod can switch between different swimming behaviours, leading to completely different flow-field patterns in self-generated surrounding flow; (ii) by exerting a time-varying propulsive force, a copepod can modulate temporally the basic flow modes to create an unsteady feeding current which manipulates precisely the trajectories of entrained food particles over a long time period; (iii) the modulation process may be energetically more efficient than exerting a constant propulsive force onto water to create a constant feeding current of a wider entrainment range. A probable reason is that the modulated unsteady flow entrains those water parcels containing food particles and leaves behind those without valuable food in them.     

Detection and capture of inert particles by calanoid copepods: the role of the feeding current

Although there is a scarcity of supporting empirical evidence,it has long been suspected that calanoid copepods use mechanoreceptionto detect the presence and location of potential prey itemsentrained in the feeding current. In this study, we documentthe first observations showing a freely swimming calanoid copepod,Skistodiaptomus oregonensis, attacking prey-sized, non-motile,inert particles entrained in the feeding current before theparticles contact the copepod's sensory appendages. Feedingcurrent geometry, fluid velocities and associated behavioursthat characterize these interactions are described. The resultsof this study show how copepod swimming behaviour, coupled witha low-velocity feeding current, not only increases copepod encounterrates with inert prey by increasing direct contact rates, butalso increases the probability of detecting and capturing remotelylocated prey that have well-developed escape responses. In turbulentregimes, a far-reaching, low-velocity feeding current shouldincrease encounter rates, but only if coupled with behavioursthat quickly minimize separation distances once prey is detected.     

Hydrodynamic interaction between two copepods: a numerical study

Numerical simulations were carried out to compute the flow fieldaround two tethered, stationary or swimming model-copepods withvaried separation distances between them and for different relativebody positions and orientations. Based on each simulated flowfield, the power expended by each copepod in generating theflow field and volumetric flux through the capture area of eachcopepod were calculated. The geometry of the flow field aroundeach copepod was visualized by tracking fluid particles to constructstream tubes. The hydrodynamic force on each copepod was calculated.Also, velocity magnitudes and deformation rates were calculatedalong a line just above the antennules of each copepod. Allthe results were compared to the counterpart results for a solitarycopepod (stationary or swimming) to evaluate the hydrodynamicinteraction between the two copepods. The calculations of thepower and volumetric flux show that no energetic benefits areavailable for two copepods in close proximity. The results ofthe stream tube and force calculations show that when two copepodsare in close proximity, the hydrodynamic interaction betweenthem distorts the geometry of the flow field around each copepodand changes the hydrodynamic force on each copepod. Two beneficialroles of the hydrodynamic interactions are suggested for copepodswarms: (1) to maintain the integrity of the swarms and (2)to separate the swarming members with large nearest neighbourdistances (usually more than five body lengths). To preventstrong hydrodynamic interactions, copepods in swarms have toavoid positions of strong interactions, such as those directlyabove or below their neighbours. The results of the velocitymagnitudes and deformation rates demonstrate that the hydrodynamicinteraction between two copepods generates the hydrodynamicsignals detectable by the setae on each copepod's antennules.Based on the threshold of Yen et al. (1992), the results showthat the detection distance between two copepods of comparablesize is about two to five body lengths. Copepods may employa simple form of pattern recognition to detect the distance,speed and direction of an approaching copepod of comparablesize.     

Chemoreception and the deformationof the active space in freely swimming copepods: a numerical study

A three-dimensional alga-tracking, chemical advection–diffusionmodel was used to calculate the deformation of the active spacesurrounding an alga entrained within the flow field around afreely swimming copepod. From the model, the advance warningtime resulting from the copepod's chemo-reception of the entrainedalga was quantified, and copepod chemoreception capability comparedfor several different swimming behaviors: hovering in the water,swimming slowly (swimming upward, swimming backward and swimmingforward), swimming fast (swimming upward, swimming backwardand swimming forward) and sinking (with the anterior pointingupward or downward). The results show that when it hovers orswims slowly, a copepod can use chemoreception to remotely detectindividual algae entrained by the flow field around itself.In contrast, a fast-swimming copepod is not able to rely onchemoreception to remotely detect individual algae. The possibilityof a free-sinking copepod using chemoreception to detect algalparticles is also indicated. It is shown that advection by thefluid motion dominates over diffusion in transporting the chemicalsignals inside the active space to the location of a copepod'schemoreceptors. The feeding current structure for a hoveringcopepod is described. It is suggested that the feeding currentstructure and re-routing or re-orienting response by a copepodin response to its antennule or other cephalic appendage inputsallow the copepod to capture the food particles that would otherwisepass outside its capture area and increase the amount of foodcaptured.     

Persistent spatial and temporal abundance patterns for the late-stage copepodites of Centropages typicus (Copepoda: Calanoida) in the US Northeast continental shelf ecosystem

The annual cycle of abundance and the monthly distributions of the copepod Centropages typicus are described for US Northeast Continental Shelf waters from samples collected on broadscale plankton surveys during 1977-87. High numbers of the copepod were captured throughout the region during the autumn months in weak south-north and onshore-offshore abundance gradients. The highest individual station densities were found near the mouth of the major estuaries and the heaviest broad-scale concentrations were usually located where bottom depth ranged from 20 to 39 m. Numbers declined throughout the ecosystem after winter arrived, less so in the southern half of the region where C.typicus abundance remained high year round in nearshore and midshelf waters between New York City and Chesapeake Bay. The copepod's abundance fell to much lower levels further north in the Georges Bank and Gulf of Maine subareas, and disappeared entirely from shelf waters in the northernmost offshore region until summer. Interannual abundance variability was substantial, but no long-term trend was detected. Analyses of samples collected from 1988 to 1996 on Georges Bank during early autumn indicate that abundance levels of C.typicus have been high here in the 1990s, completely recovered from low density values measured there in 1986 and 1987. Temperature and food availability were found to be the key factors that determine the copepod's distribution and annual abundance cycle. Mean abundance was high throughout the ecosystem where surface temperature was >9C and in regions where annual mean chlorophyll levels exceeded 1 mg m^-3. The copepod's abundance appeared to be independent from variation in water column salinity.      

Perceptive performance and feeding behavior of calanoid copepods

The goal of this study was to determine variables associatedwith calanoid feeding behavior, and thus, to improve our understandingof the basics of calanoid feeding rates. These variables includedperiods and frequency of appendage motion, rates of cell clearance,distance at which a copepod first reacts to a cell which iseventually captured, and rate of water flow through the areacovered by the motions of a copepod's feeding appendages. Theeffects of these variables on feeding rates were determinedfor copepodids and adult females of the calanoid copepod Eucalanuspileatus at phytoplankton concentrations covering the rangeencountered by this species on the south-eastern shelf of theUSA. Our results indicate that the distance at which E.pileatusperceives phytoplankton cells increases {small tilde}2-foldas food concentrations decrease from 1.0 to 0.1 mm³ l^–1.These results lead us to hypothesize that this is due to increasedsensitivity of chemosensors on the copepods' feeding appendages.This 2-fold increase in perceptive distance amounts to a near4-fold increase in perceived volume which is close to the 6-foldincrease in volume swept clear (VSC) from 1.0 to 0.1 mm³ l^–1of Thalassiosira weissflogii. We assume that the increases inVSC by planktonic copepods, when food levels are below satiation,are largely a function of the sensory performance of the individualcopepod.     

Net and gross incorporation of nitrogen by marine copepods fed on N-labelled diatoms: Methodology and trophic studies

The stable isotope of nitrogen (¹⁵N) and an appropriate three-compartment model were used in two 24-h lasting feeding experiments to trace the flow of N through the copepod Acartia discaudata and Calanus helgolandicus fed on ¹⁵N-labelled Skeletonema costatum and Thalassiosira weissflogii, respectively. Details of the labelling technique and principles of the computation of N transport rates are given. At the end of a single 24-h feeding period only about one third of the total amount of N ingested by A. discaudata was incorporated into the copepod's body N; we refer to this rate as net incorporation. Most of the N ingested was lost as ammonium (48% of total N ingested), followed by losses in the form of eggs + fecal pellets (13%) and dissolved organic N (DON, 9%). The sum of net incorporation and the latter losses is defined as gross incorporation. Net incorporation by C. helgolandicus and N losses did not vary over time during a 24 h lasting time-series feeding experiment. On average, 79% of total N ingested was actually incorporated by the copepod whereas mean N losses as ammonium, eggs + fecal pellets represented only 12 and 9%, respectively. After a 24-h feeding period only 2% of N ingested was lost as DON. Inspection of individual DON pathways showed that both A. discaudata and C. helgolandicus highly contributed to total DON production via direct excretion (79 and 64%, respectively). The remaining DON appearing in the DON pool was derived from phytoplankton via direct release and/or indirect release (copepod ‘sloppy feeding’).     

Algal toxins alter copepod feeding behavior

Using digital holographic cinematography, we quantify and compare the feeding behavior of free-swimming copepods, Acartia tonsa, on nutritional prey (Storeatula major) to that occurring during exposure to toxic and non-toxic strains of Karenia brevis and Karlodinium veneficum. These two harmful algal species produce polyketide toxins with different modes of action and potency. We distinguish between two different beating modes of the copepod's feeding appendages-a "sampling beating" that has short durations (<100 ms) and involves little fluid entrainment and a longer duration "grazing beating" that persists up to 1200 ms and generates feeding currents. The durations of both beating modes have log-normal distributions. Without prey, A. tonsa only samples the environment at low frequency. Upon introduction of non-toxic food, it increases its sampling time moderately and the grazing period substantially. On mono algal diets for either of the toxic dinoflagellates, sampling time fraction is high but the grazing is very limited. A. tonsa demonstrates aversion to both toxic algal species. In mixtures of S. major and the neurotoxin producing K. brevis, sampling and grazing diminish rapidly, presumably due to neurological effects of consuming brevetoxins while trying to feed on S. major. In contrast, on mixtures of cytotoxin producing K. veneficum, both behavioral modes persist, indicating that intake of karlotoxins does not immediately inhibit the copepod's grazing behavior. These findings add critical insight into how these algal toxins may influence the copepod's feeding behavior, and suggest how some harmful algal species may alter top-down control exerted by grazers like copepods.     

Trophic modification of food quality by heterotrophic protists: species-specific effects on copepod egg production and egg hatching

Recent literature suggests that heterotrophic protists could improve the biochemical constituents of poor quality algae for subsequent use by higher trophic levels, a phenomenon dubbed “trophic upgrading”. We conducted experiments to test trophic upgrading effects on the omnivorous calanoid copepod Acartia tonsa. Two heterotrophic protists, Oxyrrhis marina and Gymnodinium dominans, were grown on three algal species of different nutritional qualities: Dunaliella tertiolecta, Isochrysis galbana and Rhodomonas salina. In parallel incubations, the copepods were fed the heterotrophic protists or the algae that the protists grew on. We measured the ingestion rate, egg production rate and egg hatching success of the copepods in each treatment. Comparison of the egg production efficiency (EPE; egg carbon production/carbon ingestion×100%) between parallel diet treatments allowed us to calculate the trophic upgrading index (TUI; ratio of EPE-heterotrophic protist to EPE-alga). The copepods ingested similar amount, or 20-30% more algae than heterotrophic protists. Daily rations ranged 50-122% of body C per day. Copepods feeding on O. marina that grew on D. tertiolecta had higher egg production rate and egg hatching success than copepods feeding on D. tertiolecta directly, and the resultant TUI between the two treatments was 19.7. Similarly, the copepod egg production was improved by G. dominans that grew on D. tertiolecta and the resultant TUI was 23.3. TUI was near 1 between I. galbana treatment and the parallel heterotrophic protist treatments, indicating no or little trophic upgrading effects on the copepod's egg production. However, egg hatching success was significantly lower with G. dominans growing on I. galbana. O. marina that grew on R. salina induced a lower EPE relative to the alga itself, yielding a TUI of less than 1. Biochemical data showed that the heterotrophic protists contained eicosapentanoic acid (EPA) and docosahexanoic acid (DHA) even when they fed on alga that lacked these essential fatty acids, which may explain some of the observed trophic upgrading effects and species-specific differences in our experiments. However, our data also suggested that dietary EPA and DHA could not be the sole factors regulating the copepod's reproductive output, and that egg production and egg hatching appeared to have different nutritional requirements and respond differently to the copepod's diets.     

Force transmission via axial tendons in undulating fish: a dynamic analysis

Sonomicrometrics of in vivo axial strain of muscle has shown that the swimming fish body bends like a homogenous, continuous beam in all species except tuna. This simple beam-like behavior is surprising because the underlying tendon structure, muscle structure and behavior are complex. Given this incongruence, our goal was to understand the mechanical role of various myoseptal tendons. We modeled a pumpkinseed sunfish, Lepomis gibbosus, using experimentally-derived physical and mechanical attributes, swimming from rest with steady muscle activity. Axially oriented muscle-tendons, transverse and axial myoseptal tendons, as suggested by current morphological knowledge, interacted to replicate the force and moment distribution. Dynamic stiffness and damping associated with muscle activation, realistic muscle force generation, and force distribution following tendon geometry were incorporated. The vertebral column consisted of 11 rigid vertebrae connected by joints that restricted bending to the lateral plane and endowed the body with its passive viscoelasticity. In reaction to the acceleration of the body in an inviscid fluid and its internal transmission of moment via the vertebral column, the model predicted the kinematic response. Varying only tendon geometry and stiffness, four different simulations were run. Simulations with only intrasegmental tendons produced unstable axial and lateral tail forces and body motions. Only the simulation that included both intra- and intersegmental tendons, muscle-enhanced segment stiffness, and a stiffened caudal joint produced stable and large lateral and axial forces at the tail. Thus this model predicts that axial tendons function within a myomere to (1) convert axial force to moment (moment transduction), (2) transmit axial forces between adjacent myosepta (segment coupling), and, intersegmentally, to (3) distribute axial forces (force entrainment), and (4) stiffen joints in bending (flexural stiffening). The fact that all four functions are needed to produce the most realistic swimming motions suggests that axial tendons are essential to the simple beam-like behavior of fish.     

Lipids in Acanthodiaptomus denticomis during starvation and fed on three different algae

Variations in the levels of triglycerides, wax esters and polarlipids were determined in adults of the calanoid copepod Acanthodiaptomusdenticornis when freshly caught, starved or fed on the followingalgae: Anabaena spiroides, Cyclotella pseudostelligera and Pediastrumduplex. Over 7 days starvation, triglycerides and wax esterswere almost entirely used up by the copepods. Subsequent feedingover 20 days partially restored triglycerides but restored onlya relatively small fraction (<20%) of wax esters in the animals.Differences in the lipid restoration were found: the restoredtriglyceride level was higher in animals feeding on Cyclotellapseudostelligera or Pediastrum duplex than in those feedingon Anabaena spiroides. Fatty acid composition of neutral lipidswas closely linked to fatty acid composition of algae. The resultssuggest that lipid and fatty acid contents of Acanthodiaptomusdenticornis are good indices of the copepod's nutritional statusand short-term (0–20 days) feeding history.     

Implications of the feeding current structure of Euchaeta rimana, a carnivorous pelagic copepod, on the spatial orientation of their prey

Many marine planktonic organisms create water currents to entrainand capture food items. Rheotactic prey entrained within thesefeeding currents often exibit escape reactions. If the directionof escape is away from the feeding current, the prey may successfullydeter predation. If the escape is towards the center of thefeeding current, the prey will be re-entrained towards its predatorand remain at risk of predation. The direction of escape isdependent on (i) the ability of the prey to escape in a directiondifferent than its pre-escape orientation and (ii) the orientationcaused by the interaction of the prey's body with the movingfluid. In this study, the change in orientation of Acartia hudsonicanauplii as a result of entrainment within the feeding currentof Euchaeta rimana, a planktonic predatory copepod, was examined,When escaping in still water, A.hudsonica nauplii were ableto vary their pre-escape direction by only 10. This allowsonly a limited ability to escape in a direction different thantheir pre-escape orientation. Analyses of the feeding currentof E.rimana show the flow speed to be most rapid in the centralregion with an exponential decrease in speed distally. In contrast,flow vorticity is minimal in the center of the feeding currentand maximal at 1.75 mm along the antennae. As a result, thedegree of rotation of the prey towards the center of the feedingcurrent shows a strong dependency on the prey's location withinthe feeding current. The feeding current of E.rimana rotatedthe prey 14 when near the center of the flow field and up to160 when located more distal in the feeding current Since theprey's escape abilities cannot compensate for the rotation dueto the flow, this mechanism will maintain the escaping preywithin the feeding current of their predator. Therefore, thefeeding current facilitates predatory copepods in capturingprey by (i) increasing the amount of water which passes overtheir sensors and through their feeding appendages and (ii)controlling the spatial orientation of their prey prior to escape.     

Change in knee contact force with simulated change in body weight

The relationship between obesity, weight gain and progression of knee osteoarthritis is well supported, suggesting that excessive joint loading may be a mechanism responsible for cartilage deterioration. Examining the influence of weight gain on joint compressive forces is difficult, as both muscles and ground reaction forces can have a significant impact on the forces experienced during gait. While previous studies have examined the relationship between body weight and knee forces, these studies have used models that were not validated using experimental data. Therefore, the objective of this study was to evaluate the relationship between changes in body weight and changes in knee joint contact forces for an individual's gait pattern using musculoskeletal modeling that is validated against known internal compressive forces. Optimal weighting constants were determined for three subjects to generate valid predictions of knee contact forces (KCFs) using in vivo data collection with instrumented total knee arthroplasty. A total of five simulations per walking trial were generated for each subject, from 80% to 120% body weight in 10% increments, resulting in 50 total simulations. The change in peak KCF with respect to body weight was found to be constant and subject-specific, predominantly determined by the peak force during the baseline condition at 100% body weight. This relationship may be further altered by any change in kinematics or body mass distribution that may occur as a result of a change in body weight or exercise program.     

Ovigerity,selective predation,and variable diel vertical migration in Euchaeta elongata (Copepoda: Calanoida)

Summary We present a statistical analysis of a previously published (Yen, 1983) but heretofore unanalyzed data set on the vertical distributions and diel vertical migration (DVM) of adult females of the marine planktonic copepod Euchaeta elongata in Dabob Bay, Washington, USA. Non-ovigerous females were strongly migratory on all four dates sampled, residing between 75–175 m during the day and at shallower depths during the night, commonly entering the upper 50 m of the water column. In contrast, ovigerous females were non-migratory or weakly migratory, largely remaining between 100–175 m both day and night, and entering the upper 50 m of the water column only rarely. Thus non-ovigerous females always migrated much more strongly, as measured by both amplitude of migration and the proportion of animals migrating, than did ovigerous females. These results led us to hypothesize that differential susceptibility to visually orienting predators was the cause of these differences in DVM behavior in female E. elongata, and we subsequently undertook an experimental study of the feeding selectivity of the copepod's natural predator, Pacific herring (Clupea harengus pallasi). Pacific herring exhibited a highly significant preference for ovigerous over nonovigerous adult female E. elongata. The demographic consequences of variable DVM in adult female E. elongata were investigated by way of life table analyses. Results indicated that under conditions of thermal stratification of the water column there is a distinct demographic disadvantage (reduced rate of realized population growth) incurred by non-migratory or weakly migratory ovigerous females due to delayed egg development at cooler subsurface temperatures. We conclude that ovigerous female E. elongata remain at depth both day and night to avoid visually orienting predators, and that such behavior must afford the copepod a demographic advantage of no less than a 26% reduction in adult mortality to offset the demographic cost of delayed egg development.     

Feeding biology of Calanus: a new perspective

Calanus has been, for a variety of reasons, one of the most popular subjects of copepod feeding studies, and much of what we have learned from studies of Calanus has been applied to other species of copepods. Nearly all the major factors controlling feeding rate and behavior in Calanus have been known for more than three decades. These forcing functions include light, body weight, temperature, the quantity, size and quality of food, and feeding history. The relationships between these variables are better understood than they were three decades ago, but the current knowledge of them still fails to explain extraordinary variance in observations of feeding rate.I suggest that the current understanding fails because our fundamental perception is incorrect. It is generally assumed that the feeding behavior we observe is the net response to instantaneous values of a suite of functions. Past values of forcing functions may be considered a factor, but subservient to those in the present. The critical change in perspective suggested here requires that we assume the integrals of forcing functions to be more significant than their present values in regulating feeding behavior.     

Insights into the ecology and evolutionary success of crocodilians revealed through bite-force and tooth-pressure experimentation

Background

Crocodilians have dominated predatory niches at the water-land interface for over 85 million years. Like their ancestors, living species show substantial variation in their jaw proportions, dental form and body size. These differences are often assumed to reflect anatomical specialization related to feeding and niche occupation, but quantified data are scant. How these factors relate to biomechanical performance during feeding and their relevance to crocodilian evolutionary success are not known.

Methodology/Principal Findings

We measured adult bite forces and tooth pressures in all 23 extant crocodilian species and analyzed the results in ecological and phylogenetic contexts. We demonstrate that these reptiles generate the highest bite forces and tooth pressures known for any living animals. Bite forces strongly correlate with body size, and size changes are a major mechanism of feeding evolution in this group. Jaw shape demonstrates surprisingly little correlation to bite force and pressures. Bite forces can now be predicted in fossil crocodilians using the regression equations generated in this research.

Conclusions/Significance

Critical to crocodilian long-term success was the evolution of a high bite-force generating musculo-skeletal architecture. Once achieved, the relative force capacities of this system went essentially unmodified throughout subsequent diversification. Rampant changes in body size and concurrent changes in bite force served as a mechanism to allow access to differing prey types and sizes. Further access to the diversity of near-shore prey was gained primarily through changes in tooth pressure via the evolution of dental form and distributions of the teeth within the jaws. Rostral proportions changed substantially throughout crocodilian evolution, but not in correspondence with bite forces. The biomechanical and ecological ramifications of such changes need further examination.     

Danger of zooplankton feeding: the fluid signal generated by ambush-feeding copepods

Zooplankton feed in any of three ways: they generate a feeding current while hovering, cruise through the water or are ambush feeders. Each mode generates different hydrodynamic disturbances and hence exposes the grazers differently to mechanosensory predators. Ambush feeders sink slowly and therefore perform occasional upward repositioning jumps. We quantified the fluid disturbance generated by repositioning jumps in a millimetre-sized copepod (Re ∼ 40). The kick of the swimming legs generates a viscous vortex ring in the wake; another ring of similar intensity but opposite rotation is formed around the decelerating copepod. A simple analytical model, that of an impulsive point force, properly describes the observed flow field as a function of the momentum of the copepod, including the translation of the vortex and its spatial extension and temporal decay. We show that the time-averaged fluid signal and the consequent predation risk is much less for an ambush-feeding than a cruising or hovering copepod for small individuals, while the reverse is true for individuals larger than about 1 mm. This makes inefficient ambush feeding feasible in small copepods, and is consistent with the observation that ambush-feeding copepods in the ocean are all small, while larger species invariably use hovering or cruising feeding strategies.     

Functional morphology of ventral tail bending and prehensile abilities of the seahorse,Hippocampus kuda

Unlike most teleosts, the seahorse (genus Hippocampus) is able to bend its tail ventrally, uses its tail in a postural role as a grasping and holding appendage, and possesses heavy body plates instead of scales. To investigate seahorse axial bending mechanisms and the role of plating in those mechanisms, observations were made on seahorses curling their tails ventrally and holding a support and components of the mechanical system used for axial bending, including dermal plates, vertebrae, and axial muscles, were examined. Anatomical modifications involved in ventral tail bending include hypertrophy of the ventral region of the hypaxial muscle, ventrolateral attachment of the myomeres to plates, and modification of the infracarinalis posterior muscles so that they act in axial bending rather than in fin movement as has previously been hypothesized (Harder, '75) for other fishes. Modifications for prehension include the presence of fibers histochemically characterized as tonic in the median ventral muscles (the modified infracarinalis muscle) and in portions of the myomeres. Dermal plates are an important part of the force transmission system used in seahorse tail bending. They transmit forces from the hypaxial myomeres to bend the tail both laterally and ventrally. This study expands our understanding of axial bending in fishes by examining extreme modifications of the musculoskeletal system associated with the evolution of unique functional capabilities within teleosts. © 1996 Wiley-Liss, Inc.     

Work that body: fin and body movements determine herbivore feeding performance within the natural reef environment

Herbivorous fishes form a keystone component of reef ecosystems, yet the functional mechanisms underlying their feeding performance are poorly understood. In water, gravity is counter-balanced by buoyancy, hence fish are recoiled backwards after every bite they take from the substrate. To overcome this recoil and maintain contact with the algae covered substrate, fish need to generate thrust while feeding. However, the locomotory performance of reef herbivores in the context of feeding has hitherto been ignored. We used a three-dimensional high-speed video system to track mouth and body kinematics during in situ feeding strikes of fishes in the genus Zebrasoma, while synchronously recording the forces exerted on the substrate. These herbivores committed stereotypic and coordinated body and fin movements when feeding off the substrate and these movements determined algal biomass removed. Specifically, the speed of rapidly backing away from the substrate was associated with the magnitude of the pull force and the biomass of algae removed from the substrate per feeding bout. Our new framework for measuring biting performance in situ demonstrates that coordinated movements of the body and fins play a crucial role in herbivore foraging performance and may explain major axes of body and fin shape diversification across reef herbivore guilds.