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.     

Cannibalistic feeding behavior of the brackish-water copepod Sinocalanus tenellus

Cannibalistic feeding behavior of the brackish-water copepodSinocalanus tenellus was examined in the laboratory using CI-II,CIII-IV and CVI female as predators and NI-II, NIII-IV, NV-VIand CI-II as prey. In each prey-predator combination, the ingestionrate increased with increasing prey density to an asymptoticvalue. Cannibalism took place even when phytoplankton was availableas an alternative food supply. Based on a daily ration, theoptimal prey stages for CVI females, CIII-IV and CI-II are NI-VI,NI-IV and NI-II respectively. Under average, natural prey density(10 nauplii l^–1), S tenellus can achieve only a smallfraction (max 9%) of the daily minimum food requirement by cannibalisticfeeding. However, the impact of cannibalism on naupliar survivorshipcan be significant. When adult females occur at a density of10 l^–1, the mortality due to cannibalism attains 99.2%during the naupliar stages.     

Contrasting physiological responses to future ocean acidification among Arctic copepod populations

Widespread ocean acidification (OA) is modifying the chemistry of the global ocean, and the Arctic is recognized as the region where the changes will progress at the fastest rate. Moreover, Arctic species show lower capacity for cellular homeostasis and acid‐base regulation rendering them particularly vulnerable to OA. In the present study, we found physiological differences in OA response across geographically separated populations of the keystone Arctic copepod Calanus glacialis. In copepodites stage CIV, measured reaction norms of ingestion rate and metabolic rate showed severe reductions in ingestion and increased metabolic expenses in two populations from Svalbard (Kongsfjord and Billefjord) whereas no effects were observed in a population from the Disko Bay, West Greenland. At pH_T 7.87, which has been predicted for the Svalbard west coast by year 2100, these changes resulted in reductions in scope for growth of 19% in the Kongsfjord and a staggering 50% in the Billefjord. Interestingly, these effects were not observed in stage CV copepodites from any of the three locations. It seems that CVs may be more tolerant to OA perhaps due to a general physiological reorganization to meet low intracellular pH during hibernation. Needless to say, the observed changes in the CIV stage will have serious implications for the C. glacialis population health status and growth around Svalbard. However, OA tolerant populations such as the one in the Disko Bay could help to alleviate severe effects in C. glacialis as a species.     

The role of photoreception in the swarming behavior of the copepod Dioithona oculata

The copepod Dioithona oculata forms dense swarms near mangrove prop roots that are centered around shafts of light penetrating the mangrove canopy. Swarms can be created in the laboratory within light shafts created with a fiber optic light pipe. Laboratory observations of swarming behavior were recorded using video cameras, and the swimming behavior of the copepods and density of the swarms were quantified using video‐computer motion and image analysis techniques. Swarm formation results from a combination of phototactic and klino‐kinetic behavior. Dark adapted copepods initially exhibit a photophobic response to a light shaft, but become positively phototactic within 3–5 min after exposure to the light. Copepod aggregation rates under the light fit a saturation model, suggesting that copepods are attracted independently to the swarm marker. Copepods reverse their swimming direction when they encounter light intensity gradients near the edge of a light shaft, which aids in maintaining the swarm. Swarm formation can occur in the laboratory at light intensities as slow as 0.1 μM photons m^‐2 s^‐1, which is similar to light intensities at dawn when they are first observed to form in nature. Swarm formation appears to have an endogenous rhythm, as copepods will not form swarms at night under a light shaft.     

The role of the orbital cortex in feeding behavior]

It has been found that extirpation of the orbital zones of the cerebral cortex in dogs in which the structures of the limbic and striate levels of nervous integration were previously ablated, produced more profound disturbances of the higher nervous activity than the previous operations. Not only conditioned positive and inhibitory reactions were affected, but unconditioned alimentary reflexes as well, while food motivation sharply decreased. Further experiments on the operated animals and could go on, only when alimentary excitability was enhanced. A conclusion has been drawn that the orbital zones of the cerebral cortex are directly related to the achievement of an integrated alimentary behavioral reaction.     

The swimming behavior of the copepod Metridia pacfica

The swimming behavior of the copepod Metridia pacifica was studied.Animals exposed to algae showed lower average swimming speedand fewer high-speed bursts. Animals exposed to Artemia naupliialone exhibited no change in swimming behavior.     

Multilocus evidence for globally distributed cryptic species and distinct populations across ocean gyres in a mesopelagic copepod

Zooplanktonic taxa have a greater number of distinct populations and species than might be predicted based on their large population sizes and open‐ocean habitat, which lacks obvious physical barriers to dispersal and gene flow. To gain insight into the evolutionary mechanisms driving genetic diversification in zooplankton, we developed eight microsatellite markers to examine the population structure of an abundant, globally distributed mesopelagic copepod, Haloptilus longicornis, at 18 sample sites across the Atlantic and Pacific Oceans (n = 761). When comparing our microsatellite results with those of a prior study that used a mtDNA marker (mtCOII, n = 1059, 43 sample sites), we unexpectedly found evidence for the presence of a cryptic species pair. These species were globally distributed and apparently sympatric, and were separated by relatively weak genetic divergence (reciprocally monophyletic mtCOII lineages 1.6% divergent; microsatellite F_ST ranging from 0.28 to 0.88 across loci, P < 0.00001). Using both mtDNA and microsatellite data for the most common of the two species (n = 669 for microsatellites, n = 572 for mtDNA), we also found evidence for allopatric barriers to gene flow within species, with distinct populations separated by continental landmasses and equatorial waters in both the Atlantic and Pacific Ocean basins. Our study shows that oceanic barriers to gene flow can act as a mechanism promoting allopatric diversification in holoplanktonic taxa, despite the high potential dispersal abilities and pelagic habitat for these species.     

Taste perception and feeding behavior in nonhuman primates and human populations

Recent advances in taste physiology provide evidence against the traditional “western” notion that there are only four basic tastes. Each substance elicits a singular “signature” on the peripheral taste nerve, but in some cases the signals form separate clusters within the continuum of taste perceptions. We will discuss the taste abilities of nonhuman primates in terms of threshold and above-threshold responses to potential foods. As diets have evolved in species' environments, tastes have responded adaptively in order to maximize energy intake. In turn, food plants have evolved nutrients and toxins in relation to the tasting abilities of consumers. These compounds can be beneficial or harmful in various environments and at different concentrations. This cost-benefit ratio concerns all primates, including Homo sapiens populations living at subsistence level, and must be taken into account, together with psychosensory and sociocultural factors, to understand food choices.     

Preference for ethanol in feeding and oviposition in temperate and tropical populations of Drosophila melanogaster

The natural habitat of Drosophila melanogaster Meigen (Diptera: Drosophilidae) is fermenting fruits, which can be rich in ethanol. For unknown reasons, temperate populations of this cosmopolitan species have higher ethanol resistance than tropical populations. To determine whether this difference is accompanied by a parallel difference in preference for ethanol, we compared two European and two tropical African populations in feeding and oviposition preference for ethanol‐supplemented medium. Although females of all populations laid significantly more eggs on medium with ethanol than on control medium, preference of European females for ethanol increased as ethanol concentration increased from 2 to 6%, whereas that of African females decreased. In feeding tests, African females preferred control medium over medium with 4% ethanol, whereas European females showed no preference. Males of all populations strongly preferred control medium. The combination of preference for ethanol in oviposition, and avoidance or neutrality in feeding, gives evidence that adults choose breeding sites with ethanol for the benefit of larvae, rather than for their own benefit. The stronger oviposition preference for ethanol of temperate than tropical females suggests that this benefit may be more important in temperate populations. Two possible benefits of ethanol for which there is some experimental evidence are cryoprotection and protection against natural enemies.     

The role of brain somatostatin receptor 2 in the regulation of feeding and drinking behavior

Somatostatin was discovered four decades ago as hypothalamic factor inhibiting growth hormone release. Subsequently, somatostatin was found to be widely distributed throughout the brain and to exert pleiotropic actions via interaction with five somatostatin receptors (sst_1–5) that are also widely expressed throughout the brain. Interestingly, in contrast to the predominantly inhibitory actions of peripheral somatostatin, the activation of brain sst₂ signaling by intracerebroventricular injection of stable somatostatin agonists potently stimulates food intake and independently, drinking behavior in rodents. The orexigenic response involves downstream orexin-1, neuropeptide Y₁ and μ receptor signaling while the dipsogenic effect is mediated through the activation of the brain angiotensin 1 receptor. Brain sst₂ activation is part of mechanisms underlying the stimulation of feeding and more prominently water intake in the dark phase and is able to counteract the anorexic response to visceral stressors.     

Numerical study of the feeding current around a copepod

Three-dimensional, numerical simulations of the feeding current around a tethered copepod were performed using a finite-volume code. The copepod's body shape was modeled to resemble Euchaeta norvegica, and was represented by a curvilinear body-fitted coordinate system. In the simulations, the appendages that generate the feeding current were replaced by a distribution of forces acting on the water adjacent and ventrally to the body. First, the accuracy of the code was verified by simulating two viscous, zero-Reynolds-number flows for which analytical solutions are available. Then, simulations with realistic body shape and Reynolds numbers were carried out. The main features of the computed feeding current were compared with observations from Yen and Strickler (Invert. Biol., 115, 191-205, 1996), and good agreement was obtained. The entrainment region, as visualized by tracking particles in the feeding current and by plotting the resulting stream-tube, is quite large. The result can be used to quantify how the copepod takes advantage of the feeding current to trap the algal particles in its capture area. The configuration of the feeding current near to the body surface of the copepod is controlled by how the copepod forces the feeding current and by the copepod's morphology. These parameters were varied and their effects studied in a systematic manner. Specifically, by comparing various spatial distributions of the same amount of total force, it was shown that a distributed force dissipates less energy (and increases the entrainment rate) than a concentrated force, it is thus energetically more desirable. Variations of the copepod's body shape and of the distribution of forces showed little effect on the far field of the feeding current, and therefore do not appear to affect the detectability by other mechano-receptional organisms. The length scale of the influence field of the feeding current was shown to be anisotropic in three directions, extending 5-7 mm above or ventrally to the copepod, <1 mm dorsally to the copepod and >1 cm down from the abdomen. The results also suggest that the net reaction force on the copepod from the feeding current is of the same order of magnitude as the excess weight of the copepod, but is not sufficient to balance the excess weight completely.      

Nature's role in sustaining economic development

In this paper, I formalize the idea of sustainable development in terms of intergenerational well-being. I then sketch an argument that has recently been put forward formally to demonstrate that intergenerational well-being increases over time if and only if a comprehensive measure of wealth per capita increases. The measure of wealth includes not only manufactured capital, knowledge and human capital (education and health), but also natural capital (e.g. ecosystems). I show that a country''s comprehensive wealth per capita can decline even while gross domestic product (GDP) per capita increases and the UN Human Development Index records an improvement. I then use some rough and ready data from the world''s poorest countries and regions to show that during the period 1970–2000 wealth per capita declined in South Asia and sub-Saharan Africa, even though the Human Development Index (HDI) showed an improvement everywhere and GDP per capita increased in all places (except in sub-Saharan Africa, where there was a slight decline). I conclude that, as none of the development indicators currently in use is able to reveal whether development has been, or is expected to be, sustainable, national statistical offices and international organizations should now routinely estimate the (comprehensive) wealth of nations.     

Effects of diet and starvation on digestive enzyme activity and feeding behavior of the marine copepod Calanus pacificus

In this paper we look at the response of Calanus pacificus overa period of several weeks to both starvation and diet as reflectedby survival rates, feeding rates and digestive enzyme activities.Phytoplankton species used as food were the diatom Thalassiosiraweissflogii, two dinoflagellates, Gymnodinium simplex and Scrippsiellatrochoidea, and the chlorophyte Dunaliella tertiolecta. In general,C.pacificus appeared to be resistant to starvation for periodsup to 1 week. Survival remained high for at least 6 days ofstarvation, while maximum feeding rates remained constant forseveral days and then declined by 50% after a week of starvation.When initial digestive enzyme activities were low, they didnot change appreciably during the experiments. However, activitylevels dropped rapidly during starvation experiments in whichthey were initially high. Other evidence indicates that starvationmay induce changes in the digestive system before it affectsingestion rates or survival. In general, long-term changes indigestive enzyme activity under different dietary regimes weregradual. An exception to this pattern was S. trochoidea, whosepresence in the diet induced a precipitous, though reversible,decline in digestive enzyme activity, a phenomena possibly relatedto the toxic reaction to this species reported by other researchers.A positive induction of digestive enzymes occurred over a time-scaleof 1–5 days in response to a change in diet. The absenceof laminarin in the diet of C.pacificus for nearly 3 weeks (whenfeeding on G.simplex), during which time laminarinase activitydeclined by {small tilde}50%, did not adversely affect subsequentassimilation of a laminarin-rich food (T. weissflogii).     

A rapid analysis of copepod feeding using FlowCAM

This study addressed the usefulness and reliability of usinga new plankton image analyzer, FlowCAM, for rapid analysis ofcopepod feeding by comparison with the conventional microscopicanalysis. We carried out bottle incubation experiments withtwo copepod species in the Oyashio region and analyzed the preyabundance prior to and after the incubation with a FlowCAM.From the volume-specific fluorescence intensity of particles,the FlowCAM successfully distinguished between zooplankton andphytoplankton and allowed an adequate evaluation of the copepodfeeding on zooplankton and phytoplankton. The analysis timefor one plankton sample was about 10 min, which was less thanone-tenth of the time required for microscopic enumeration.The FlowCAM is considered to be an efficient tool for rapidanalysis of copepod feeding particularly in studies of omnivory.     

Swimming and feeding behaviour of the planktonic copepod Clausocalanus furcatus

The objective of this study was to quantify motion and feeding behaviour of the small calanoid Clausocalanus furcatus, which is a common and abundant species in oligotrophic environments. Adult females were video recorded at 60 fields s^-1 using video equipment which allowed us to follow each free-swimming individual continuously in 3 1 vessels. At 20C under dark conditions, C.furcatus moved continuously along convoluted small loops at a mean speed of 10 mm s^-1 (corresponding to 10 body lengths s^-1). This motion was occasionally interrupted by sudden somersaulting performed at very high speed (up to 17 mm s^-1). The copepods only occasionally sank. There was no evidence that C.furcatus created feeding currents. Dinoflagellate cells offered as food appeared to be perceived by direct encounter at high speed, being instantaneously captured and ingested or rejected. The capture rates were correlated with the frequency of somersaulting. By moving fast and by associating a high turning rate with a high frequency of crossing the previous tracks, C.furcatus searched 22-26% of the explored volumes. The motion and feeding behaviour of C.furcatus show that the foraging tactic of this species is to explore small volumes of water rapidly. This strategy appears, so far, to be unique among small planktonic copepods.      

The role of the cerebral ganglia in the organization of feeding behavior in the pteropod mollusk Clione limacina]

The role of cerebral ganglia in switching from free swimming over to hunting behaviour in Clione limacina (Phipps) was studied. The cell bodies of central neurones giving processes to the sensory and motor nerves were mapped by retrograde axonal transport of Co2+ ions, and behavioural reactions under the extracellular electric stimulation of different ganglia areas were investigated. Comparison of morphological and behavioural data suggests main role of cerebral ganglia neurones in switching of behaviour from free swimming over to any of the three possible states: hunting (including food uptake), active or passive defensive behaviour.     

The components of feeding behavior in rotifers

Feeding behavior of a rotifer can be broken into two classes of activities: the rate of successful search and the handling process. The former consists of the following components: Perceptual field (in planktonic rotifers the area of the corona), swimming rate, and attack rate. The second class consists of capture rate, handling time, rejection rate, ingestion, digestion, and assimilation. All evidence indicates that the perceptual field cannot be varied by the rotifer. Swimming rate is variable and under rotifer control, but does not appear to vary with degree of starvation. Attack rate is also under control of the rotifer, at least in the genus Asplanchna. Capture rates vary with the species of food item from zero to 100%. Handling times are longer than one would expect, as are rejection times. Digestion and assimilation appear to vary inversely with rate of ingestion. There is some suggestion in the literature that feeding behavior on very small particles differs from that on larger ones.     

Aggregation of helminths: the role of feeding behavior of fish hosts

Individual Arctic charr (Salvelinus alpinus) from Fjellfr?svatn, northern Norway, could be categorized by their stomach contents as zooplanktivores or benthivores. Feeding specialization among these fish was evident from negative correlations between helminths transmitted by pelagic copepods (Diphyllobothrium dendriticum and D. ditremum) and those transmitted by the benthic amphipod Gammarus lacustris (Cystidicola farionis and Cyathocephalus truncatus). Occurrences of parasite species acquired from the same types of invertebrate were positively correlated in the fish. Strong relationships among habitat use, diet, and helminth infections among the Arctic charr indicated persistent foraging patterns involving long-term habitat use and feeding specialization. The distribution of all parasite species was highly aggregated in the fish samples, measured by the exponent k of the fitted negative binomial distributions (range: 0.5-7.5) and the variance-to-mean ratios (s2/mean, range: 5-85). Charr specializing on either copepods or Gammarus predominantly contributed to high-intensity class intervals within the overall frequency distributions of the corresponding parasite species. Such fish had low infection intensities of helminths transmitted by other prey organisms. The detailed analyses of the parasite frequency distributions for fish with different habitat or feeding preferences evidently show how heterogeneity in trophic behavior contributes strongly to the commonly observed aggregation of helminths among hosts under natural conditions.     

The role of termites in tropical forestry

Summary Tropical forestry is influenced by termites in widely differing ways.Large mounds constructed byMacrotermes in Africa and Asia are ecological factors which influence the natural regeneration of forests and the vegetation patterns of savanna woodlands and grasslands.Economic losses in timber caused by termite attack on mature trees are restricted to those areas in Australia, Asia and Central America where particular species ofCoptotermes are present.Afforestation withEucalyptus in less humid areas of Africa, India and South America has been greatly facilitated by the use of organochlorine insecticides to protect seedlings and saplings from attack by ground-dwelling termites.

---

Zusammenfassung Für die tropische Forstwirtschaft ist eine Reihe von Termitenarten in unterschiedlicher Weise bedeutsam.Dis grossen Termitenhügel vonMacrotermes sind in Afrika und Asien ökologisch von grosser Wichtigkeit, indem sie die Regeneration des Waldes und die Vegetationsverteilung in Trockensavannen und Grasland beeinflussen.Wirtschaftliche Holzschäden durch Termitenangriff auf stehende Waldbäume beschränken sich in Australien, Asien und Zentralamerika auf solche Gebiete, in denen best-immteCoptotermes-Arten vorkommen.Die Aufforstung mit Eucalyptusarten in Afrika, Indien und Südamerika ist wesentlich erleichtert worden, seitdem man dazu übergegangen ist, die Sämlinge und Jungbäumchen durch verschiedene organischsynthetische-Boden-Insektizide zu schützen.

     

Role of host genetics and heat‐tolerant algal symbionts in sustaining populations of the endangered coral Orbicella faveolata in the Florida Keys with ocean warming

Identifying which factors lead to coral bleaching resistance is a priority given the global decline of coral reefs with ocean warming. During the second year of back‐to‐back bleaching events in the Florida Keys in 2014 and 2015, we characterized key environmental and biological factors associated with bleaching resilience in the threatened reef‐building coral Orbicella faveolata. Ten reefs (five inshore, five offshore, 179 corals total) were sampled during bleaching (September 2015) and recovery (May 2016). Corals were genotyped with 2bRAD and profiled for algal symbiont abundance and type. O. faveolata at the inshore sites, despite higher temperatures, demonstrated significantly higher bleaching resistance and better recovery compared to offshore. The thermotolerant Durusdinium trenchii (formerly Symbiondinium trenchii) was the dominant endosymbiont type region‐wide during initial (78.0% of corals sampled) and final (77.2%) sampling; >90% of the nonbleached corals were dominated by D. trenchii. 2bRAD host genotyping found no genetic structure among reefs, but inshore sites showed a high level of clonality. While none of the measured environmental parameters were correlated with bleaching, 71% of variation in bleaching resistance and 73% of variation in the proportion of D. trenchii was attributable to differences between genets, highlighting the leading role of genetics in shaping natural bleaching patterns. Notably, D. trenchii was rarely dominant in O. faveolata from the Florida Keys in previous studies, even during bleaching. The region‐wide high abundance of D. trenchii was likely driven by repeated bleaching associated with the two warmest years on record for the Florida Keys (2014 and 2015). On inshore reefs in the Upper Florida Keys, O. faveolata was most abundant, had the highest bleaching resistance, and contained the most corals dominated by D. trenchii, illustrating a causal link between heat tolerance and ecosystem resilience with global change.