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.     

Evolutionary implications of swimming behaviour in meiobenthic copepods

Body morphology is said to be the all important factor in determining swimming prowess in copepods. Fusion and differentiation of the body (tagmosis) is coupled with advance into the pelagic realm of the Gymnoplea and is thought, by the provision of a rigid thoracic tagma, to promote swimming efficiency. Thus pelagic copepods are believed to be secondarily derived from bottom dwelling predecessors. Experimental evidence is presented to show that the majority of bottom dwelling harpacticoid families, including the most primitive and the most advanced, have representatives that undergo active sustained swimming movements. Such a widespread occurrence is indicative of a conservative evolutionary trait. This primitive behaviour is linked to precopulatory association which takes place necessarily in the water column; it is a feature retained by representatives of all copepod orders. The implication of cephalic appendage vibration (feeding currents) is the essential feature in the swimming success of the Gymnoplea; planktonic efficiency in these is suggested to have evolved coincident with, rather than because of increased tagmosis.     

Perception of inert particles by calanoid copepods: behavioral observations and a numerical model

High-resolution video showed freely swimming Diaptomus sicilisattacking and capturing inert 5O µm polystyrene beadsthat were outside the influence of the copepod feeding current.The beads were frequently more than half a body length awayand were attacked after the ‘bow wake’ of the movingcopepod displaced the bead away from the copepod. To investigatethe hypothesis that deformation of streamlines around the copepodand its first antennae stimulated the attack response, a finiteelement numerical model was constructed. The model describedthe fluid interactions between a large object approaching asmaller object in a laminar flow at Reynolds number 5, whichis characteristic of the fluid regime experienced by foragingcopepods. The model revealed that fluid velocity fluctuationsand streamline deformations arose in the region between thetwo objects as separation distance between the objects decreased.The video observations and the model results support the hypothesesthat chemoreception is not required for the detection and captureof large phytoplankton cells [Vanderploeg et ai, in Hughes,R.N. (ed.), Behavioral Mechanisms of Food Selection. NATO ASISeries G20, 1990; DeMott and Watson, /. Plankton Res., 13, 1205-1222,1991], and that swimming behavior plays an integral role inprey detection. ⁴Present address: Academy of Natural Sciences Estuarine ResearchCenter, 10545 Mackall Road, St Leonard, MD 20685, USA     

Chemoreception in an arctic amphipod crustacean: A field study

Traps baited with meat or fish were set through the ice in 5 m of water near Prudhoe Bay, Alaska. They were monitored by divers and surface observers. Over 99% of the animals attracted to the bait were the lysianassid amphipod Boeckosimus (= Onisimus) affinis (Hansen). Up to 20000 individuals were collected during each 24-h trapping period. Animals moved to the trap only from a downcurrent direction, and from a distance of up to at least 30 m away. It is suggested that chemoreception is the method used to find bait: animals swam into the current after receiving the stimulus and required a constant presence of stimulus at some threshold level to continue moving to the bait.     

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.     

The flow field around a freely swimming copepod in steady motion. Part I: Theoretical analysis

The three-dimensional flow field around a free-swimming copepodin steady motion was studiedtheoretically. This study was basedon coupling the Navier–Stokes equations with the dynamicequations for an idealized body of a copepod. To allow analyticalsolutions to the flow field, three simplifications were made:(a) to simulate the effect of the beating movement of the cephalicappendages, a force-field was added to the Navier–Stokesequations, (b) to linearize the problem, Stokes flow was used,and (c) to simplify the morphologies of the copepods, a sphericalbody shape was assumed. Analytical solutions were derived forfive steady motions: (1) hovering, (2) sinking, (3) upwardsswimming, (4) backwards swimming and (5) forwards swimming.The results show that thegeometry of the flow field around afreely swimming copepod varies significantly with the differentswimming behaviours. When a copepod hovers in the water, orswims very slowly, it generates a wide, cone-shaped flow field.In contrast, when a copepod sinks, or swims fast, the flow geometryis not cone-shaped, but cylindrical, narrow and long. Theseresults are consistent with published observations on live copepods.It is shown that the differences in the flow geometry with thedifferent swimming behaviours are due to the relative importancebetween the two factors in generating the flow field: the copepod'sswimming motion and the requirement to counterbalance the copepod'sexcess weight. The results also highlight the importance ofconsidering freely swimming copepods as self-propelled ratherthan as towed bodies. ‘Self-propelled’ means a freelyswimming copepod must gain thrust from the surrounding waterin order to counterbalance the drag force by water and its excessweight. Regardless of swimming behaviours and velocities, thefar-field velocity field decays to that of the velocity fieldgenerated by a point force of magnitude equal to the copepod'sexcess weight in an infinite domain. On the other hand, usingthe towed body model yields a flow field with much differentfar- and near-field flow characteristics. Hence, the towed bodymodel is inherently unable to reproduce fundamental characteristicsof the flow field around a freely swimming copepod.     

Chemoreception: identifying friends and foes

The vomeronasal organ detects chemical cues that trigger sexual, aggressive and defensive behaviors. An in situ hybridization analysis has identified the specificities of nearly a hundred VNO receptors and elucidated the logic by which they encode these cues.     

Feeding and digestive activity of cyclopoid copepods in active diapause

Three different methods were used to compare the feeding patterns of active cyclopoids and cyclopoids in active diapause from different habitats. First, we offered ¹⁴C- labeled algal detritus to stage III, stage IV (two different physiological groups), and adult females of Acanthocyclops vernalis from temporary water-bodies of the Volga Delta. Second, we compared the functional responses of active and actively diapausing cyclopoids preying on copepod nauplii. Stage III, stage IV (various physiological groups), stage V, adult males and females of Acanthocyclops viridis, Cyclops kolensis, C. abyssorum sevani from the Volga Delta, Lake Sevan (Armenia), and Mozhaisk Reservoir (Moscow District) were tested. Finally, we examined the activity of digestive enzymes (proteases) of CIV of C. kolensis from Mozhaisk Reservoir at different temperatures and temperature acclimation regimes. All three approaches used have led to similar conclusions. Actively diapausing cyclopoids were capable of feeding on both plant and animal prey; however, their feeding rates and digestive enzyme activity were 2–8 times lower than those of active cyclopoids of similar size. At the initial stage of active diapause the feeding rates were comparable with those of active cyclopoids then declined to a low level during approximately two weeks. Transfer of cyclopoids undergoing winter diapause from ambient temperature (4 °C) to warmer water (20 °C) resulted in dramatic increase of proteases activity only if the acclimation period was relatively long (two weeks).     

Evidence for biorhythmicity from the preoptic nucleus of the rainbow trout Salmo gairdnerii under freely swimming conditions--an electrophysiological study

This study describes a technique which allows continuous recording of MUA (Multiple Unit Activity), from the NPO (Preoptic Nucleus), DAP (Dorsal Aortic blood Pressure) and ECG (Electrocardiogram) in freely swimming rainbow trout. From the 21 trout tested, six trout (29%) clearly showed rhythmic patterns of MUA during the five post-operative days (D2-D6). The mean length of rhythmic MUA was about 18 hr (range 6-33 hr) among the six trout during the recording days. Periodic MUA occurred approximately eight times/hr and lasted about 2 min. The maximal frequency of discharges was 20-30 spikes per sec. No change occurred in the mean level of blood pressure from the first operative day to the following post-operative days, where rhythmic MUA appeared or reappeared. These results demonstrate the existence of biorhythmicity within the NPO of freely swimming trout and suggest parallel oscillations in neurohormones secretion.     

Cannibalistic behaviour of rock-pool copepods: An experimental approach for space, food and kinship

Studies on cannibalism in harpacticoid copepods are restricted to predation on naupliar larvae in rock-pool harpacticoids of the genus Tigriopus. An earlier experimental study on the Mediterranean copepod Tigriopus fulvus indicated that females recognized their own larvae and preferentially preyed on nauplii other than their own. In a series of laboratory experiments, we tested if there were differences in naupliar predation as a function of crowding, food level and sex in Tigriopus brevicornis and T. fulvus. Results show that cannibalism was restricted to the first larval stages (N1 and N2). Both food availability and adult density significantly affected the predation rate. Contrary to earlier suggestions, adult males also preyed on the nauplii. We found no evidence that adults spare their own offspring, for neither T. fulvus nor T. brevicornis.This is in accordance with what one would expect for species having the life history characteristics of Tigriopus, i.e.: multiple broods and large number of offspring. Earlier results indicating parental care in Tigriopus must be taken with caution.     

Chemoreception in hydra: specific binding of glutathione to a membrane fraction.

Specific binding of reduced [35S]glutathione (GSH) was measured using a crude membrane fraction of Hydra vulgaris (attenuata). The specific binding shows both rapid displaceable and nondisplaceable components. Rapid displaceable binding accounted for 20% of the total specific binding. Data from saturation binding experiments indicates half maximal total specific binding occurs at 2 microM GSH which is similar to reported EC50 values from behavioral experiments. Calcium is required for displaceable binding of GSH to the putative receptor. Oxidized glutathione (GSSG), an antagonist of the GSH-activated feeding response, and S-methylglutathione (GSM), an agonist of the feeding response, inhibit the binding of radiolabeled GSH to the putative receptor. Glutamate, a putative competitive antagonist of the GSH-activated feeding response in hydra, does not inhibit the specific binding of radiolabeled GSH to the receptor and must therefore block the feeding response by a mechanism other than competitive inhibition.     

Chemoreception and feeding behaviour in a caterpillar: towards a model of brain functioning in insects

The quantitative relationship between chemoreceptor activity during food intake and total amount of food ingested per unit of time has been studied in last instar larvae of Pieris brassicae. Data on sensory responses and feeding intensity, derived from a study by Blom (1978), show a strong correlation between receptor activity during 1 sec periods of stimulation and food intake during 24 h. It is concluded that nerve impulses from chemoreceptors, which signal the presence of different phagostimulants, are summated algebraically in the central nervous system. Impulses from feeding deterrent receptors counteract the effects of phagostimulants. One impulse from a deterrent receptor neutralizes 2.5 impulses from phagostimulant receptors. Based on the relationships observed a simple model of a feeding centre in the central nervous system has been constructed (Fig. 8).     

Duetting in space: a radio-telemetry study of the black-bellied wren

In many birds, individuals 'answer' the songs of their pair-mates to produce vocal 'duets'. One hypothesized function of song answering is that it prevents extra-pair birds from intruding into the duetting pair's territory to obtain copulations or usurp one of the pair-mates. In this capacity, answering may signal that the pair-mates are close together, and so are prepared to defend against such an intrusion. Another functional hypothesis states that answering helps pair-mates maintain contact, and so predicts that a bird is more likely to approach its mate after a duet than after a solo song. I used radio-telemetry to monitor the distance between mated black-bellied wrens (Pheugopedius fasciatoventris). I found that birds of both sexes were more likely to answer their mate's song when the mate was close, and that maximum duet length was negatively related to the distance between pair-mates. Furthermore, song answering positively affected the likelihood of one pair-mate approaching the other after a song. In a significant majority of the approaches after duet songs, the answering bird approached the initiator. I conclude that in the black-bellied wren, (i) the occurrence and duration of vocal duets covary with physical closeness and (ii) contact maintenance is a secondary function of duet participation.     

AChemS: the beginning. Association for Chemoreception Sciences

This paper unfolds the events, the people and the times that led up to thefounding of AChemS and fashioned its character during its early formativeyears. It describes the path over which AChemS came, going from theoriginal assertions and denials for the need of such an organization to itslater inception and nascent development. This narration highlights suchtopics as the debate over the need for AChemS, the role of National ScienceFoundation in the founding of AChemS, the derivation of the Association'sname, the choice of Sarasota and the Hyatt House as the meeting site, thegeneration of the programs for the early annual meetings, the adoption ofthe bylaws, the process of incorporation and tax deferment, and the birthof the Givaudan Lectureship. Most emphatically highlighted, however, is theenthusiasm, commitment and hard work that the members of the chemosensoryresearch community displayed in bringing AChemS to fruition.     

Strategies for sperm chemotaxis in the siphonophores and ascidians: a numerical simulation study

Chemotactic swimming behaviors of spermatozoa toward an egg have been reported in various species. The strategies underlying these behaviors, however, are poorly understood. We focused on two types of chemotaxis, one in the siphonophores and the second in the ascidians, and then proposed two models based on experimental data. Both models assumed that the radius of the path curvature of a swimming spermatozoon depends on [Ca(2+)](i), the intracellular calcium concentration. The chemotaxis in the siphonophores could be simulated in a model that assumes that [Ca(2+)](i) depends on the local concentration of the attractant in the vicinity of the spermatozoon and that a substantial time period is required for the clearance of transient high [Ca(2+)](i). In the case of ascidians, trajectories similar to those in experiments could be adequately simulated by a variant of this model that assumes that [Ca(2+)](i) depends on the time derivative of the attractant concentration. The properties of these strategies and future problems are discussed in relation to these models.