In situ diel variation in gut pigment contents of Ceriodaphnia sp. in stratification and destratification periods

The short-term, in situ diel grazing of Ceriodaphnia sp. duringperiods of stratification and mixing was investigated usingthe technique of fluorimetric analysis of the gut pigments.There were considerable seasonal differences in feeding behaviourIn mixing, when the concentration of chlorophyll a in the watercolumn was high and Ceriodaphnia abundance was low, gut pigmentcontents showed clear diel variation patterns, probably dueto diel variations of the high values of feeding activity observedin the 24 hour cycle The maximum values were found at dawn.On the other hand, no diel variations in gut pigment were observedduring periods of stratification and while the amounts of pigmentsin the water and in the gut were very low, species abundancewas high. Taking into account the ambient conditions, the authorsdiscuss the possibility that the change of feeding of the Ceriodaphniasp. observed when the environment changed from a mixing periodto one of stratification represents a change from herbivorousto detritivorous behavior.     

Feeding, excretion and egg production by individuals and populations of the marine, planktonic copepods, Acartia spp. and Centropages furcatus

Diel variations in vertical distribution, gut pigment content,ammonium excretion and egg production were investigated foradult females of Acartia erythraea and A.pacifica in the verticallymixed Inland Sea of Japan and Centropages furcatus in the stratified,neritic Gulf of Mexico. Gut pigment content and egg productionrate were maximal at night and ammonium excretion was maximalduring the daytime. Neither A.erythraea nor A.pacifica adultfemales showed an apparent diel migration, but the former werehighly concentrated in the surface layer during the afternoon.In contrast, C.furcatus adult females showed a clear diel migration,residing immediately above the bottom during the daytime andbeing concentrated between 10 and 25 m depth during the nighttime.Individual-based data on gut content and excretion and egg productionrates were combined with vertical-distribution data to calculatepopulation values. In the Inland Sea of Japan, the resultantpattern for Acartia spp. reflected the diel variation in physiologicalrates and even distribution of adult females, except for theafternoon, surface aggregation of A.erythraea. In the Gulf ofMexico, the pattern for C.furcatus reflected largely the dielvariation in each rate process and the heterogeneous distributionof adult females in the water column. Elevated nocturnal feedingactivity of these copepods may be due to an endogenous rhythm.The daytime maximum in ammonium excretion and night-time maximumin egg production rate indicated approximate half-day and daytime lags, respectively, after the intake of food until itsconversion into dissolved excreta and released eggs.     

Diel variations in gut pigment content, diel vertical migration and estimates of grazing impact for copepods in the southern Benguela upwelling region in October 1987

The gut fluorescence technique was used to estimate ingestionand filtration rates of the adult female copepods Paracalanusparvus, Cenlropages brachiatus and Calanus austrails, and copepoditestages 3, 4 and 5 of C.australis in the southern Benguela upwellingregion. During the study period chlorophyll concentrations withinthe upper 20 m of the water column were high, 5 µg I^–1in mid-shelf waters and 15–30 µg I^–1 in innershelf waters. Copepod gut pigment content was low and constantduring the day then increased sharply during the first 2 h aftersunset. Gut pigment content was 2–6 times higher duringthe night compared with daytime values. Small non-migratingcopepods (Paracalanus parvus) showed the smallest diel differencein gut pigment content and large migrating copepods (Centropagesbrachiatus and Calanus australis) the largest difference. Eggproduction rates were 20 and 50% of maximum at the mid-shelfand inner shelf stations respectively, suggesting food-limitation.Comparison of ingestion rates calculated from egg productiondata with ingestion rates calculated from gut pigment data suggestedthat the copepods were feeding omnivorously at the inner shelfstations but herbivorously at the mid-shelf stations. Assumingthat all of the phytoplankton was available as food, the nearshorecopepod assemblage grazed {small tilde}1% of the standing cropeach day, and the mid-shelf assemblage grazed 5% day^–1.Because of errors and uncertainties associated with the gutfluorescence technique, the feeding impact could be underestimatedby 2–4-fold. We discuss several approaches which couldlead to more precise estimates of feeding rates. ³Present address: Marine Sciences, SUNY, Stony Brook, NY, 11794-5000,USA     

Zooplankton-phytoplankton interactions in a eutrophic lake

Enclosure experiments were made in a cyanobacteria dominatedlake (Lake Rotongaio) to assess the impact of zooplankton (>150µm) grazing on algal growth rates and determine the effectof diel and vertical changes in zooplankton grazing intensityand nutrient (NH₄-N) regeneration upon abundance of phytoplankton.The filamentous cyanobacterium Anabaena minutissima var. attenuataand diatom Cyclotella meneghiniana showed a negative linearchange in abundance with a gradient in zooplankton grazing intensity.Phytoflagellates were not grazed and showed a positive linearchange in abundance with increasing zooplankton biomass. Theseeffects, as well as shortening of filament length of Anabaena,were caused by raptorial feeding by the alanoid copepod Boeckellapropinqua which dominated the zooplankton. Phytoplankton growthwas not stimulated by addition of nutrients, suggesting nutrientregeneration was not important. Diel and vertical changes infeeding and NH₄-N regeneration rates were measured in Marchand June 1988. Diel differences were more pronounced in Marchwhen the water column was stratified. Specific feeding rateswere more important than vertical changes in zooplankton biomassin determining community grazing rates in March, but in Junewhen the water column was mixed, vertical distribution of zooplanktonbiomass was important. Zooplankton grazing was an importantloss process for phytoplankton in the lower part of the epilimnionin Lake Rotongaio.     

Influence of phytoplankton composition and stratification degree on gut pigment content of Ceriodaphnia sp. at dawn

Very short-term feeding activity of the cladoceran Ceriodaphniasp. was investigated in situ in a eutrophic reservoir in thesouth of Spain, using fluorimetric analysis of the gut pigmentcontent in periods when the water column was relatively mixedor strongly stratified. The results obtained in the mixed watercolumn showed a clear increase in gut pigment content at dawn,a period sampled with high frequency. The accumulation of thecladoceran at the depth of maximum concentration of phytoplankton,and the high gut pigment concentration in cladocerans at thatdepth just after dawn, suggested active feeding of Ceriodaphniaon phytoplankton at that time. During stratification, the abundanceof Ceriodaphnia was higher, but the gut pigment contents werevery low and they did not reflect any clear feeding patterns,with either time or depth. Changes in phytoplankton concentrationand composition between the relatively mixed and the stratifiedwater column suggest a shift in feeding activity from herbivorousto.     

The use of gut fluorescence to estimate grazing by oceanic salps

The fluorometric measurement of chlorophyll and its degradationproducts in the guts of salps has been used as a means of calculatingin situ grazing rates. Salps are particularly suitable for thismethod because they are non-selective filter-feeders large enoughto be individually collected by divers. Analysis of the pigmentin the gut has shown that chlorophyll degradation products canbe used as tracers to estimate in situ ingestion. Filteringrates are calcuLated from the amount of pigment, the gut clearancetime and the concentration of pigment in the water. For Pegeaconfoederata and Salpa maxima, gut-pigment rates are comparedwith rates measured in laboratory particle-clearance experiments.Salpa cylindrica has proved impossible to maintain in the laboratoryand the gut-pigment results are the first feeding data for thisspecies. Filtering rates for all species examined are high,on the order of liters per hour. Variability in the amount ofpigment found in S. cylindrica suggests this species, unlikethe others, may feed sporadically. The gut-pigment method permitsmeasurements of filtration and ingestion by herbivores on analmost instantaneous basis and can be a useful tool for assessinggrazing impact in field situations.     

Feeding behaviour and migrations in a natural population of the copepod Acartia tonsa

This paper deals with the variations on feeding activities and diel migrations of Acartia tonsa Dana, the dominant copepod species in Berre lagoon (west Mediterranean French coasts).A 27 hour in situ study was carried out during June 1989, at a station located in the south west of the lagoon. Vertical profiles of salinity, temperature and dissolved oxygen taken each 12 hours showed a stratification of the water column in two distinct layers: (1) a superficial layer with higher temperature, moderate salinity, and high oxygen concentrations; (2) a colder, more saline and almost anoxic deep layer. In situ chlorophyll a measurements were made at –1, –3 and –5 m; concentrations were relatively homogeneous through the water column during the whole sampling period.Zooplankton samples were collected every 3 hours with a 200 µm mesh net, in three strata (0–2 m, 2–4 m, 4–6 m). A complete dominance of A. tonsa was observed in the zooplankton community. Our results point out clearly a nocturnal migration, with individuals concentrating in both superficial layers; thus an unimodal migratory pattern can be inferred. Gut flourescence measured following the Mackas & Bohrer technique (1976), showed higher values during night time, and values for females were the highest with wider day—night variations. Similar results were found in laboratory experiments with copepods fed with a culture of Dunaliella tertiolecta.Gut evacuation rate was measured in two laboratory experiments either mixing or separating males and females. Evacuation rate was 17.92 and 27.25 min for males and females respectively.Phytoplankton daily ration for A. tonsa calculated by gut flourescence and gut evacuation rate was particularly low, for it represents only 10% of the individual carbon weight.Moreover, grazing impact on phytoplankton is very restricted, it represents less than 1% of the daily phytoplankton stock.     

Fluorescence analysis of zooplankton gut contents and an investigation of diel feeding patterns

A method is described for measurement of the in situ feeding rate of planktonic herbivores. Freshly caught Zooplankton are washed with filtered sea water and frozen. The gut contents are later analysed using acetone extracts of the entire sample. Fluorescence peaks due to ingested chlorophyll are easy to quantify for sample sizes of 20–200 animals. This method obviates some of the problems, such as confinement and previous feeding history, associated with extrapolation from laboratory experiments to field conditions. The method is illustrated by an investigation of diel feeding patterns of marine copepods. Initial results indicate the complexity of these patterns, with either a single night peak or dawn and dusk peaks for adult copepods. Diel variations in feeding activity appear to occur in addition to, but closely interacting with, the diel vertical migrations of the animals.     

Feeding rhythm and vertical migration of the euphausiid Euphausia pacifica in coastal waters of north-eastern Japan during fall

We investigated the feeding rhythm and diel vertical migrationof Euphausia pacifica in coastal waters of north-eastern Japanin September 2000. The euphausiids arrived at the surface earlyin the night and sank just after satiation. Gut pigment contentsand stomach fullness were higher in the food-rich surface layerat night than in the food-poor deep layer during the day. Feedingactivity of E. pacifica during daytime was low even if foodorganisms were abundant in the ambient water in the top 50 m.Therefore, the feeding rhythm of E. pacifica is explained bydecreased feeding activity during the day rather than by reducedfood availability.     

Pigment destruction by Calanus pacificus: impact on the estimation of water column fluxes

Measurements of gut pigment made prior to fecal pellet productionin Calanus pacificus females and CV copepodites suggest that(i) chlorophyll a and/or its pheopigment derivatives are degradedinto molecules that are not detectable by the standard fluorometrictechnique; and (ii) the percentage of ingested chlorophyll awhich degrades into fluorometrically undetected molecules isnot constant. Thus, measurements of chlorophyll and pheopigmenta in the guts of zooplankton can only yield minimum estimatesof in situ grazing rates. Estimates of the vertical flux ofprimary particulates based on chlorophyll and pheopigment abudgets may also be underestimated.     

Sexual dimorphism, activity budget and synchrony in groups of sheep

The activity budget hypothesis has been proposed to explain the social segregation commonly observed in ungulate populations. This hypothesis suggests that differences in body size – i.e. between dimorphic males and females – may account for differences in activity budget. In particular, if females spend more time grazing and less time resting than males, activity synchrony would be reduced. Increased costs of maintaining synchrony despite differences in activity budget would facilitate group fragmentation and instability of mixed-sex groups. In this paper two prerequisites of the activity budget hypothesis were tested: (1) that males should spend less time feeding and more time resting than females in single-sex groups and (2) that lower activity synchrony should be observed in mixed-sex compared to single-sex groups. The activity budget and synchrony in mixed and single-sex groups of merino sheep (Ovis aries) of different sizes (2, 4, 6, 8 individuals) were measured in three contiguous 491-m² arenas located in a natural pasture. Three same-size groups, one of each category, were observed simultaneously. We found no sexual differences in the time spent inactive and active (i.e. grazing, standing, moving, interacting). Males spent significantly more time grazing and less time standing than females. These differences disappeared when yearling males were omitted from the group. Males and females had similar bite and step rates. Sheep of both sexes spent less time resting and more time grazing and moving and had lower bite rates when in mixed-sex groups than when in single-sex groups. The synchrony among visually isolated groups was near zero, indicating that they changed activities independently. On the contrary, within-group synchrony was high; however it was higher in single-sex groups, in particular for males, than in mixed-sex groups. Our results suggest that differences in activity budget and synchrony alone are insufficient to explain social segregation.     

Is fecundity modified by crowding in the copepod Centropages typicus?

Experiments were conducted to test the effect of overcrowdingon egg production rates in the copepod Centropages typicus.Fertility was not significantly reduced by the presence of otherconspecifics when the volume of water was the same for femalesincubated alone or in the presence of other females and males.There was no evidence of chemical cues affecting egg productionwhen females were kept in large (1 female in 100 ml sea water[SW] or small (1 female in 25 ml SW) volumes of water, withor without partitions, and in the presence or absence of foodor males. Also, spawning rate was not reduced when females wereincubated in flow-through experimental chambers which alloweddiffusion of pheromones between females incubated alone or inhigh densities (100 ind. in 50 ml SW). Only when females wereplaced at unnaturally high densities (1 female ml^–1) wasegg production significantly reduced, probably due to physicaldisturbance and increased number of collisions rather than chemicalcommunication among individuals. The results show that chemicalcues do not affect spawning behaviour when population densitiesare close to field conditions. By contrast, overcrowding inducedby swarming or aggregation may potentially limit reproductivesuccess due to reduced feeding, and production and cannibalismof eggs.     

Diel patterns of zooplankton grazing in a shallow lake

A diel survey of in situ species-specific zooplankton clearancerates (with radioactively labelled Chlorella cells) was conductedin the shallow lake of Cr?teil characterized by small-sizedplanktonic forms (algae<10µm and zooplankton <1.3mm). Experiments were performed every 4h at two depths (1 and4m). Power functions relating individual filtering rates tobody length were established for the three most abundant cladoceransand for calanoids (nauplii being included in this feeding group),for each depth and time. No filtering penodicity was observedin Ceriodaphnia spp., adults and copepodites of Eudiaptomusgracilis and Eurytemora velox, and nauplii. On the contrary,clear nocturnal filtering peaks were obtained for Daphnia spp.and for Diaphanosoma brachyurum, these being more pronouncedfor the larger individuals at 1 m deep. The observed diel periodicitycannot be explained by variations in physico-chemical parametersor food concentration alone. The ecological significance ofthis phenomenon in the polymictic lake of Cr?teil is discussedin the light of previously published data and the hypothesesaccounting for it.     

Diel activity of adult pikeperch <Emphasis Type="Italic">Sander lucioperca</Emphasis> (L.) in a drainage canal in the Mediterranean basin during spring

Pikeperch (Sander lucioperca L.) is a broadly distributed fish species in Europe but little is known about its ecology in the southern part of its distribution area in warm climatic conditions. The aim of this study was to analyse pikeperch rate of movement and to assess whether it displayed a diel pattern related to temperature. Thus acoustic telemetry was used to track adult pikeperch in a drainage canal located in south of France. The survey was carried out in spring, during the spawning period. The results showed that females were more active than males. This is in accordance with previous data on the nest guardian behaviour of the males. For both genders, the activity rates increased during the study period as water temperature rose. Males and females displayed the same diel activity with a maximum at dusk, thereby confirming many indirect observations. Nevertheless, inter-individuals variations were observed. Thus, these results on diel activity are rather a general trend than a strict rule and suggest the involvement of other factors than light intensity in the control of diel activity. This diel rhythm is positively correlated to water temperature for females. Pikeperch activity may be the result of a trade-off between physiological requirements of temperature and light, satisfaction of energy needs and avoidance of predators.     

Egg production of individual copepods of Acartia grani Sars from coastal waters: seasonal and diel variability

Seasonal and diel fecundity patterns of adult Acartia granifemales from a coastal area of Malaga Bay (south of Spain) werestudied between spring and autumn 1990. Copepod egg productionwas measured in situ by short-term (     

In situ measurement of filtering rates of the salp, Thalia democratica, on phytoplankton and bacteria

Salps were gently captured in a perspex chamber by a SCUBA diver,who then injected a suspension of ¹⁴C-labeled phytoplanktonand ³H-labeled bacteria into the chamber. After 0.5 h incubationin situ, filtering rates were estimated from incorporation ofthe two isotopes, and expressed as a function of each salp'ssize. Both bacteria and phytoplankton were grazed by Thalia,the latter at higher rates. Weight-specific grazing rates increasedwith increasing size of salp. Estimated hourly rations rangedfrom 1% of bodily C for small Thalia feeding only on phytoplanktonto 8% of bodily C for large Thalia feeding on all particulateorganic C. The method gave repeatable results for Thalia, butwas unsatisfactory for the pteropod, Cavolinia. ¹Present address: Institute of Marine Resources, A-018, Universityof California, San Diego, Scripps Institution of Oceanography,La Jolla, CA 92093, USA.     

Diel patterns of grazing by pigmented nanoflagellates on <Emphasis Type="Italic">Synechococcus</Emphasis> spp. in the coastal ecosystem of subtropical western Pacific

Natural populations of planktonic Synechococcus spp. exhibit diel variations in abundance and frequency of dividing cells (FDC) during the warm seasons (>25°C) in coastal water of the western subtropical Pacific ocean. We hypothesized that differences in grazing rates during the day/dark cycle were a major cause of the observed diel variations in Synechococcus spp. abundance. We used fluorescently labeled particles (FLP) as tracers of feeding on Synechococcus spp. in June and August 2007. Our results showed that FDC of Synechococcus spp. were highest at dusk (50%) and lowest (<10%) between midnight and early morning. Synechococcus spp. had three abundance phases, accrual (I), peak (II), and diminished (III). Moreover, FDC values gradually increased in phase I, declined to the lowest values during phase II, and remained at low levels throughout phase III. Our results strongly indicated that pigmented nanoflagellate (PNF) grazing was the underlying biological factor regulating diel variations in Synechococcus spp. abundance. And, the rate of PNF ingestion peaked during the night on smaller non-dividing cells following a peak of Synechococcus spp. abundance in phase II. These findings provide evidence that diel variations in ingestion rates are affected by non-dividing cells of Synechococcus spp. and imply that the impacts of PNF grazing on Synechococcus spp. is based on food selectivity by size.     

Evidence of selective feeding on larvaceans by the pelagic copepod Candacia bipinnata (Calanoida: Candaciidae)

Gut contents of Candacia bipinnata collected from the InlandSea of Japan and the neighbouring sea revealed that these pelagiccalanoid copepods feed selectively on larvaceans.     

Gut clearance and pigment destruction in a herbivorous copepod, Acartia tonsa, and the determination of in situ grazing rates

Gut clearance rates of starving and continuously feeding Acartiatonsa were estimated. During the initial 30 min the rates weresimilar (0.045 and 0.048 min^–1, respectively; 14°C)but thereafter starving animals expelled the remains of theirgut contents at half the rate (0.019 min^–1) of fed ones(0.048). Pigment destruction was estimated by (i) incubationexperiments over 3–4 days, (ii) silica to pigment ratioin algae and faeces and (iii) by gut filling experiments. Theincubations showed that 8% of the ingested pigments were destroyedto nonfluorescent residues during gut passage. The silica topigment ratio method gave an average of 11 % (1 –24) destructionand gut-filling experiments showed no systematic differencebetween ingestion measured as gut filling rate (fluorescence)and particle reduction. ¹Present address: Kristineberg Marine Biological Station, S-45034 Fiskebäckskil, Sweden     

Activity Pattern of Polecats Mustela putorius L. in Relation to Food Habits and Prey Activity

The activity pattern of polecats Mustela putorius L. was studied by radiotracking five males and four females in the wetlands of western France from Jul. 1988 to Mar. 1992. Polecats were nocturnal throughout the year with no differences between males and females. The diel activity remained moderate (31 %). Seasonal food habits were characterized by the exploitation of anurans in spring, while rodents were preyed upon from summer to winter. Seasonal changes in the activity rhythm of polecats appeared synchronized with the activity of the main prey. It is suggested that the activity patterns exhibited in different periods of the year are a result of changes in feeding tactics.