On assessment of prey ingestion by copepods

The consumption of photosynthetic and heterotrophic cells byan abundant calanoid copepod species feeding on natural planktoncommunities was quantified with a state-of-the-art image-analysissystem. Late copepodid stages of Eucalanus pileatus did notingest bactena, small photosynthetic and heterotrophic nanoplankton,or the abundant Ceratium spp. in quantifiable amounts Althoughdiatoms were by far the most abundant cells (in terms of POC1^–1), the copepods ingested a higher percentage of ciliatesin relation to their abundance than of diatoms and small heterotrophicdinoflagellates in the first experiment, and ingested a higherpercentage of dinoflagel lates and ciliates compared to diatomsin the second experiment Heterotrophic cells sufficiently largeto be captured were repeatedly preferred by E.pileatus overautotrophs of similar or larger size. More over, among the cellswhich could be individually perceived by this calanoid, largerones were not pre ferred over smaller cells, implying that someaspect of food quality can be as significant as prey size. Theseresults support the notion (e.g. Kleppel, Mar. Ecol Prog. Ser.,99, s183–195, 1993) that feeding by copepods will be underestimatedif ingestion of heterotrophic food organisms is not quantified.While the proposed microscope-based method is comparativelyslow (     

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.     

On Predator Selection against Abnormalities of Movement A Test of an Hypothesis1

If debilitated prey items can be caught more efficiently than healthy ones, there is a selective advantage for predators to detect any weakness in their prey. This study investigates whether three-spined sticklebacks (Gastcrosteus aculeatus L.) accustomed to feeding on one or the other of two types of waterfleas differing in their swimming pattern capture the unfamiliar type when given a choice. Such sequential oddity selection might serve as a mechanism for the detection of sick prey.     

Effects of concentration on the feeding of a marine copepod in algal monocultures and mixtures

High-speed microcinematography was used to examine the feedingbehavior of the marine copepod Eucalanus elongatus in a rangeof concentrations of algal monocultures and mixtures. Two celltypes were offered, the 13-µm diatom Thalassiosira weissflogii,which is primarily accumulated passively by low amplitude flappingof the second maxillae, and the 20 450-µm diatom Rhizosoleniaalata, which is actively captured by detection of and orientedresponse to individual cells. E. elongatus rapidly switchedback and forth between these two capture modes in mixtures ofboth diatoms, and flapped the second maxillae at low amplitudesregardless of the absolute or relative abundance of small andlarge cells. However, copepods in both monocultures and mixturesaltered the duration and/or rate of flapping of the feedingappendages with changes in algal concentration, with maximumactivity levels occurring at intermediate concentrations. Themarked reduction in feeding motions observed at the lowest algalconcentrations supports results from traditional grazing studiesand optimal foraging models, and may conserve energy duringprolonged perods of low food availability in continental slopewaters.     

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.     

Particle capture by Favella sp. (Ciliata, Tintinnina)

Particle capture by the tintinnid ciliate, Favella sp., wasinvestigated with high-speed video microscopy. Experiments withmicrospheres and algal cells indicate that within the size range1–19 p.m. behavioral responses are partly responsiblefor the differential capture of large over small particles.For example, 4 µm microspheres or Isochrysis [spheiicaldiameter (ESD) - 4.5 µ are only captured if they are encounteredin the inner zones of the encounter area. Ten micrometer microspheresor Heterocapsa(14–19 – are captured with a greaterefficiency than smaller particles from the inner zones, butcan also be captured from the outer zone of the encounter area.Capture of particles from the outer zone is associated withcilialy reversal. Microsp1ieres and algal cells of similar sizeare captured in a similar manner.     

Variability due to feeding activity of individual copepods

This paper addresses (the inter- and intraindividual) variabilityin release rates of fecal pellets of early copepodids (Cll)and adult females of the calanoid copepod Eucalanus pileatus.The intervals between releases of fecal pellets of tetheredindividuals were used as a measure of feeding activity. Earlycopepodids represented animals with a known, and adult femalesthose with an unknown, and probably highly variable history.Food concentration had a significant effect on the frequencyof pellet release by adult females, but not by CII. There wereno significant differences in release intervals among femalesat the lowest food level, but significant differences at middleand highest food concentrations. These differences could bepartly attributed to different history (seasonally). The coefficientof variation (CV) as a measure of variability, rarely exceeded30% for females and was always <30% for CII. For most individuals,the CV was <20%. The range in the frequency of pellet releasewas almost always less than factor of two. Our data indicatethat in pellet release frequency, variability in feeding behaviorwithin and between individual E.pilealus is rather low comparedto data on feeding rates, and is attributed largely to the appliedprocedures.     

Remote detection of algae by copepods: responses to algal size, odors and motility

Selective feeding on large algae by copepods involves remotedetection of individual particles and subsequent active captureresponses In this study we use radiotracer experiments to quantifythe clearance rates of five coexisting freshwater copepods andto investigate the relative merits of the chemoreception andmechanoreception hypotheses of remote detection Tropocyclopsand three diaptomid copepods exhibited relatively high clearancerates when feeding on low concentrations of large algae, suggestingthat most previous studies with freshwater copepods have underestimatedmaximal clearance rates and the degree of size selectivity.All five species of copepods exihibited strong selection foran intermediate-sized flagellate (25 µm Cartena) or alarge-sized nonmotile alga (80 µ.m Pediastrum) over asmall-sized flagellate (6 µ.m Chlamydomonas). The weight-specificclearance rate for Tropocylops prasmus feeding on motile Cartena(271 ml mg^–1 h^–1) was about twice that of threediaptomid copepods and more than an order-of-magnitude higherthan the estimate for Epischura lacustris feeding on its preferredalga, Pediastrum Assuming that distance chemoreception (‘smell’)is important in remote detection, we predicted that the additionof high concentrations of ‘algal odors’ would obscureany chemical gradients emanating from individual algal cellsand would thereby hinder the remote detection and active captureof large algae Contrary to this hypothesis, the addition ofamino acids, sucrose, and algal extracts had no effect on theclearance rates and selectivity of Diaptomus birgei. These results,together with recent cinematographic studies (Vanderploeg etal.,1990), suggest that mechanoreception is the primary mechanismfor the remote detection of large particles by diaptomid copepods.A raptorial cyclopoid, Tropocyclops prasinus, exhibited strongpreferences for motile algae, whereas a suspension-feeding calanoid,D birgei, did not select between motile and nonmotile cells.Motility appears to be an important factor in algal detectionfor small cyclopoid copepods but not for suspension-feedingdiaptomids     

Vertical zooplankton distribution on the northeastern Florida shelf and its relation to temperature and food abundance

The vertical distribution of abundant zooplankton on the northeastern(NE) Florida shelf was measured and related to depth, temperatureand concentration of particulate matter. High particle concentrationswere observed in low-temperature near-bottom intrusions of upwelledGulf Stream water. Patterns of vertical distribution differedamong the abundant zooplankton taxa. Abundances of the calanoidsEucalanus pileatus and Temora turbinata, late copepodids andadults of the cyclopoid Oncaea sp., the cladcceran Penilia avirostrisand larvae of the cephalochordate Branchiostoma sp. generallyincreased with increasing depth. E. pileatus, T. turbinata,Oncaea sp. and P. avirostris increased with increasing concentrationof paniculate matter. Abundances of young specimens of the tunicateOikopleura, the calanoid Centropages furcatus and during oneperiod, the cyclopoid Oithona sp. were greatest in the uppermixed layer and the thermocline. The abundance of Eucalanus,T. turbinata and the late copepodids/adults of Oncaea showeda significantly positive conelation to the abundance of paniculatematter.     

The use and misuse of public information by foraging red crossbills

Group foragers may assess patch quality more efficiently bypaying attention to the sampling behavior of group members foragingin the same patch (i.e., using "public information"). To determinewhether red crossbills (Loxia curvirostra) use public informationto aid their patch departure decisions, we conducted experimentsthat compared the sampling behavior of crossbills foraging ona two-patch system (one patch was always empty, one patch containingseeds) when alone, in pairs, and in flocks of three. When foragingalone, crossbills departed from empty patches in a way thatwas qualitatively consistent with energy maximization. We foundevidence for the use of public information when crossbills werepaired with two flock mates, but not when paired with one flockmate. When foraging with two flock mates, crossbills sampledapproximately half the number of cones on the empty patch beforedeparting as compared to when solitary. Furthermore, as expected ifpublic information is used, the variance in both the numberof cones and time spent on the empty patch decreased when crossbillsforaged with two flock mates as compared to when alone. Althoughhigh frequencies of scrounging reduce the availability of publicinformation, scrounging is usually uncommon in crossbills, apparentlybecause they exploit divisible patches. Consequently, publicinformation is likely to be important to crossbills in the wild.We also show that feeding performance is greatly diminishedwhen the feeding performances of flock mates differ. This providesa mechanism that will favor assortative grouping by phenotypewhen phenotypes affect feeding performance, which may in turnpromote speciation in some groups of animals.     

家蚕味觉电生理反应的个体差异

为探讨家蚕Bombyx mori人工饲料饲养发育不齐的生理原因,从同一蚕品种中挑选出对人工饲料摄食性不同的个体,用电生理学方法测定了下颚瘤状体味觉感器对4种代表性物质（蔗糖、肌醇、大豆粉提取物和柠檬酸）的电生理反应。结果表明,栓锥感器Ss-Ⅰ对蔗糖等促食物质的反应以及栓锥感器Ss-Ⅱ对大豆粉提取物等阻食物质的反应,均存在明显的个体差异。在临界浓度下,低摄食性个体的放电脉冲频率显著高于高摄食性个体。说明低摄食性蚕的味觉反应比高摄食性蚕敏感。     

Resistance to blue-green algal toxins by Bosmina longirostris

Bosmina longirostris was resistant to strains of Microcysrtsaeruginosa and Anabaena flosaquae previously reported to havelethal toxic effects on cladocerans. These blue-green algaewere of poor nutritional value to B.longirostris; survivorshipwas increased when fed these algae but reproduction was negligible.Results of feeding selectivity experiments showed that B.longirostrisdid not avoid consuming these blue-green algae, indicating thatthe mechanism for resistance must be post-ingestion. These resultssuggest that, unlike the other cladocerans tested, B.longirostriscould potentially coexist with toxic blue-green algal blooms.     

栖息地质量对两种网蛱蝶集合种群结构和分布的影响

在河北省赤城县研究了栖息地质量对大网蛱蝶Melitaea phoebe和金堇蛱蝶Euphydryas aurinia两种网蛱蝶集合种群结构和分布的影响。这两种网蛱蝶在约10 km_²的区域内共存,成虫期的蜜源植物几乎相同,大网蛱蝶的发生峰期比金堇蛱蝶晚约一个月,两者只有不到一周左右的时间重叠。大网蛱蝶和金堇蛱蝶幼虫的寄主植物分别是: 祁州漏芦（菊科）和华北蓝盆花（川续断科）。蜜源植物的丰度与两种网蛱蝶的局域种群大小呈正相关;祁州漏芦的密度对大网蛱蝶的局域种群大小影响很大,金堇蛱蝶的局域种群大小则与其寄主植物华北蓝盆花的高度正相关;斑块内平均植被高度与两种网蛱蝶的局域种群大小均呈正相关,植物多样性、植物均匀性和植被盖度均与金堇蛱蝶的局域种群大小负相关,与大网蛱蝶的关系不大。同时分析了其他因子如斑块的坡向、坡度等的影响。主要结论是：1）幼虫寄主植物的不同和成蝶飞行峰期的分离允许两种网蛱蝶在这样一个小的斑块区域内共存;2）蜜源是重要的限制因子,并且受气候随机性的影响很大,蜜源的波动可以很好地解释网蛱蝶集合种群在年度间的动态变化;3）大网蛱蝶和金堇蛱蝶的飞行、食物搜寻能力的不同以及各自寄主植物的生物学特性、空间分布的不同决定了它们具有不同的集合种群结构： 金堇蛱蝶是经典的集合种群,大网蛱蝶是源-汇集合种群;4）斑块质量和昆虫行为共同决定了两种网蛱蝶的集合种群结构和分布。     

Nitrogen regeneration by the subtropical marine copepod Eucalanus pileatus

Ammonium release rates by individual marine copepods (Eucalanuspileatus) were examined kinetically over successive 10-min intervalsafter the animals were exposed to three concentrations of thediatom Thalassiosira fluviatilis as food. Food concentrationsspanned those expected in the natural environment. "Well-fed"(3 mm³ T. fluviatilis l^–1) copepods released ammoniumsignificantly (p<0.05) faster than those fed zero or lowconcentrations of food, but differences were not dramatic. Meanexcretion rates (± SE) for animals removed from foodfor 18 –22 h [30 ± 6 nmol NH₄(mg ash free dry weight)^–1h ^–1] were 60% of those for "well-fed" animals [49 ±8 nmol NH₄ (mg ash free dry weight)^–1 h^–1]. Continuedhigh release rates after extended periods without food suggestthat E. pileatus must feed frequently in nature to maintainan adequate nitrogen balance. Depending on food concentration,animals released ammonium at rates ranging from 40 to > 100%of their N ingestion rates over the previous 18–22 h.Long-term (4–6 h) kinetic excretion experiments with "well-fed"animals indicated that, on average, release rates decreasedwith time after food removal, but patterns varied among individualcopepods. In addition to ammonium release, o-phthalaldehydereactive amino acid nitrogen was occasionally released by E.pileatus in large "spurt events" lasting from 20 to 60 min. ¹GLERL Contribution No. 283     

Endosymbiosis in Hydra and the Evolution of Internal Defense System

Certain species of Chlorella have exploited an intracellularhabitat and occur naturally as cytobionts in Hydra viridissima.The algae evoke phagocytosis by Hydra digestive cells and aresequestered in individual phagosomes that migrate to the baseof the host cells and resist fusion with lysosomes. The abilityto resist digestion is closely correlated with release of extracellularcarbohydrate (maltose) by the algae. The established populationof algae grows at an average rate equal to or greater than thatof the host and a constant population density is maintained.The host regulates algal population density by expelling ordigesting excess algae, or by controlling algal cell division.The control mechanism is unknown but can be breached by additionof inorganic ions to the Hydra culture medium with the resultthat the algae overgrow the Hydra. The Hydra-Chlorella symbiosis is probably mutually beneficial,but conditions such as prolonged darkness (with or without feeding)can reduce the competitive fitness of the host since this conditionresults in heterotrophy by the algae at the expense of selectedhost substrates. The evolution of selective permeability toorganic substrates is a major feature of the successful colonizationof the intracellular habitat by symbiotic Chlorella.     

Top-down impact by copepods on ciliate numbers and persistence depends on copepod and ciliate species composition

Experiments with the copepods Acartia clausi, Centropages hamatusand Pseudocalanus sp. were performed to assess the species-specificeffect of these copepods on the development of monospecificalgae (Nephroselmis pyriformis) and ciliate communities (Strombidiumvestitum, Strombidium conicum, Strombidium sp. and Lohmanniellaoviformis). It was hypothesized that potentially switching copepodslike A. clausi will stabilize the algal community by switchingbetween ciliate and algal food, in contrast to copepods withstereotypic filter feeding behaviour (Pseudocalanus sp.). Intreatments with Pseudocalanus sp. and C. hamatus, all ciliatespecies were wiped out in 2 days, resulting in blooms of N.pyriformis. In treatments with A. clausi, two of the ciliatespecies were able to persist, but the combined ciliate and copepodcommunity was not able to control the algal bloom. Ciliatesbecame abundant in control treatments without copepods, butonly S. vestitum and S. conicum seemed able to establish grazingcontrol. Hence, when evaluating the role of ciliates in foodwebs, their actual numbers and species composition should betaken into account. Likewise, the species composition of copepodsmay be crucial; these experiments demonstrate that small filterfeeding copepods may have tremendous impact on ciliate numbers.     

Prey Capture in Actinopterygian Fishes: A Review of Suction Feeding Motor Patterns with New Evidence from an Elopomorph Fish, Megalops atlanticus

Suction feeding is recognized as the dominant mode of aquaticprey capture in fishes. While much work has been done identifyingmotor pattern variations of this behavior among diverse groupsof actinopterygian fishes, many ray-finned groups are stillnot represented. Further, the substantial amount of inherentvariation in electromyography makes much of the pioneering workof suction feeding motor patterns in several basal groups insufficientfor evolutionary comparisons. Robust evolutionary comparisonshave identified conserved qualitative traits in the order ofmuscle activation during suction feeding (jaw opening > buccalcavity expansion > jaw closing). However, quantitative traitsof suction motor patterns (i.e., burst durations and relativeonset times) have changed over evolutionary time among actinopterygianfishes. Finally, new motor pattern evidence is presented froma previously neglected group, the Elopomorpha. The results suggestthat future investigations of the muscles influencing lateralexpansion of the mouth cavity and head anatomy may provide valuablenew insights into the evolution of suction feeding motor patternsin ray-finned fishes. In addition, the evidence illustratesthe value of comprehensive EMG surveys of cranial muscle activitiesduring suction feeding behavior.     

Filtering rates on natural bacteria by Daphnia longispina and Eodiaptomus japonicus in Lake Biwa

Filtering rates on [³H]thymidine-labelled natural unattachedbacteria and that on [¹⁴C]bicarbonate-labelled natural planktonwhich pass through the 25 µm-mesh-size screen were measuredfor Daphnia longispina and Eodiaptomus japonicus in Lake Biwa.Errors associated with the radioisotope technique, i.e the lossof labels after feeding trials and the self-absorption of thebeta emittance of ³H, were checked and corrected for the calculationof the filtering rates. It was suggested that Daphnia collectsbacteria efficiently, although the efficiency is somewhat variabledepending on food particle composition (i.e. presence and absenceof larger particles) and feeding condition (i.e. animal densityand physical disturbance). By contrast, copepodites of Eodiaptomuswere suggested to be less efficient bacteria feeders. Food resourceexploitation strategies of these two co-existing zooplanktersare discussed.     

Selective feeding by protozoa: model and experimental behaviors and their consequences for population stability

Selective feeding by zooplankton can have profound consequencesfor the stability of grazer and prey populations, as demonstratedby the behavior of plankton dynamics models. We present an analyticalapproach—calculation of prey ratio trajectories—thatreveals unambiguously whether selective feeding behavior isstabilizing (i.e. provides a refuge for preferred prey speciesat low prey concentrations) or destabilizing (i.e. results inelimination of prey populations). Prey ratio trajectories werecalculated for three modeled selective feeding behaviors. Constantselection was consistently destabilizing, while selection behaviorsthat changed inresponse to either prey ratio or prey abundancecould be stabilizing. Batch culture experiments with four protozoangrazer species (three ciliates, one heterotrophic dinoftagellate)demonstrated that protozoa fed selectively in every case, weaklypreferring the larger of the two algal species offered. Stabilizingselection was observed only in the experiment with Favella sp.,however, meaning that only this species altered its selectionbehavior in response to changing experimental conditions. Becauseprotozoa are the major grazers of phytoplankton in many planktonicsystems, our findings indicate that the use of selective feedingbehaviors to stabilize plankton dynamics models needs carefulevaluation. The modeling and graphical techniques presentedhereare a tool for linking further exploration of selective feedingbehaviors with the development of planktondynamics models.     

Laboratory studies on the larval growth and development of Nyctiphanes capensis (Euphausiacea)

Larvae of Nyctiphanes capensis Hansen were reared in the laboratoryunder different trophic conditions (i.e., algal, animal andmixed diets) from the second calyptopis stage to the first juvenilestage. It is demonstrated that diet is important in determininggrowth rates and development of the larvae. The highest growthrates were achieved on diets of Artemia nauplii mixed with eitherthe flagellate Tetraselmis or Pseudodiaptomus nauplii. Poorerdietary conditions such as with the diatom Phaeodactylum yieldedslower growth rates where additional moults were needed to completefurcilia development. Under favourable trophic conditions theintermoult period was 4–6 days and was not age dependent.Three dominant pathways of pleopod development were found inhealthy individuals.