Size-specific swimming behavior of Daphnia pulex

Body length affects several aspects of the behavior of quietlyswimming Daphnia pulex Swimming and sinking rates were measuredat 0.033 s intervals during the ‘hops’ characteristicof Daphnia swimming behavior Larger animals swim faster, covermore distance, and produce more powerful swimming strokes. LargerDaphnia also sink faster, but the sinking rate scales as lengthto the 0 58 power, far lower than the power of 2 00 predictedby Stokes Law considerations. The number of hops s^–1 wasindependent of body size, although a theoretical analysis predictshopping rate (antennal beat frequency) should increase as bodylength squared. Turning behavior, measured as the ratio of displacementto total distance, during 5 s, is also independent of body sizeIndependence of several parameters of body motion and body sizeimplies that factors other than simple mechanics affect Daphniaswimming behavior     

Seasonal analysis of Daphnia pulicaria swimming behavior

Our study documents individual swimming behavior of Daphnia pulicaria over a yearly cycle in a temperate lake. We collected D. pulicaria, a common freshwater zooplankton, from Lake Mendota on 10 dates between July 1994 and June 1995 from two depths, 2 m and 10 m. The Daphnia were rushed to the laboratory and video-taped as they swam in lake water under lake-ambient temperature and light conditions. Five-second swimming tracks of individual Daphnia were filmed and digitized using a motion analysis system. We measured average turning angle, swimming speed and sinking rate for each track. D. pulicaria swimming behavior varied over the annual cycle. We found significant differences in turning angle between depths and among months. Sinking rate and swimming speed were significantly different among months but not depths. Sinking rate and swimming speed were not significantly correlated with water temperature. Our results were contrary to Stokes' Law predictions, in that D. pulicaria had the slowest sinking speed in June, not in the winter when water temperatures were lowest and viscosity was highest. Body length was significantly correlated with all three swimming variables. We also studied swimming behavior in clonal populations of D. pulicaria in different concentrations of the alga, Chlamydomonas reinhardtii. D. pulicaria did not change swimming speed, turning angle or sinking rate over a range of food concentrations. Finally, swimming behavior in a D. pulicaria clone, tested at two temperatures in the laboratory, confirmed the results from our seasonal study; Daphnia did not sink as predicted by changes in viscosity.     

Interactive effects of fish kairomone and light on Daphnia escape behavior

We investigated the effects of fish kairomone and light intensity on the inducibility and effectiveness of escape behavior in four clones of the water flea Daphnia from different habitats. To characterize and determine the effectiveness of their escape responses, individuals were observed: (i) escaping from the hydrodynamic disturbances of a simulated predator (a small sphere dropped from above); (ii) being preyed upon by small fish (Poecelia reticulata); (iii) responding to encounters with conspecifics in crowded conditions. The simulated predation experiments revealed that when exposed to fish kairomone for 48 h, two Daphnia pulicaria clones, but not two hybrid clones, became about twice as sensitive to fluid disturbances when tested in the light, relative to no-kairomone and dark treatments. When tested in the dark, kairomone had no effect on sensitivity in any clone. All four clones had an all-or-none escape response, in which the strength of the response, as measured by escape distance and speed, was constant regardless of treatment. In the guppy predation trials, kairomone-treated D.pulicaria escaped significantly more often from guppies, in both bright- and dim-light conditions. In dim light, similar to natural lighting conditions, regardless of kairomone, all but the most weakly escaping clone were able to elude attacking guppies in a significant proportion of attacks. Finally, kairomone had no effect on the number of escapes performed by crowded individuals in response to the comparatively weak, non-threatening signals created by other Daphnia, indicating that the kairomone-induced alertness in the D.pulicaria clones did not make them 'excessively' sensitive. The results suggest that Daphnia escape behavior is under complex and efficient environmental regulation, and may play a significant role in aquatic trophic relationships.      

The interactive effects of temperature,food level and maternal phenotype on offspring size in Daphnia magna

Invertebrate offspring are usually larger in colder environments. To test for possible effects of covariates (e.g. maternal phenotype and feeding conditions) on this pattern, we performed a laboratory experiment to look at the effect of temperature on newborn weight in the planktonic crustacean Daphnia magna. Three tempèratures (12°C, 16°C and 22°C) and two food levels (10,000 cells ml^–1 and 150,000 cells ml^–1) were used, and offspring were examined from the first five clutches of mothers that had been maintained under the constant experimental conditions for three generations. Preliminary analysis suggested that newborn weight was significantly affected by temperature although patterns in the data were not clear cut. In addition, the covariates mother weight and clutch size were positively and negatively correlated with newborn weight, respectively; and later clutches tended to contain heavier offspring. Therefore, in an effort to control for the effects of the covariates, repeated-measures analysis of covariance was performed using ratio values of newborn weight/mother weight (relative newborn weight) as the dependent variable, clutch size as the covariate and clutch number as the repeated measures term. Now, temperature as a main effect in an ANCOVA model did not significantly influence relative newborn weight. The repeatedmeasure term clutch number also became nonsignificant, indicating that when differences in mother weight due to age were accounted for there were no overall differences in relative newborn weight between clutches from a particular mother. Temperature effects on relative newborn weight were only significant as part of interaction terms with food concentration and with clutch number. Thus there were different weight responses to temperature within food levels, and between clutch numbers within food levels. Under the low-food conditions newborn were heaviest at 16°C, lightest at 12°C and intermediate at 22°C. Conversely, under the high-food condition newborn were lightest at 16°C, heaviest at 12°C and again intermediate at 22°C. However, newborn tended to be heavier under the low food condition, and food concentration was highly significant as a main effect. Mother growth rate showed no significant relationship with newborn weight. It is concluded that direct temperature effects on relative newborn weight are marginal and nonsignificant. Temperature effects through interactions with food concentration and clutch number are important determinants of newborn weight, but relatively speaking account for only a small proportion of observed variance in newborn weight (25%), compared with the direct effect of food concentration (67%).     

Filter mesh size and food particle uptake by Daphnia

Food size selection of four Daphnia, species (D. magna, D. hyalina, D. galeata, D. pulicaria) was investigated using spherical plastic beads as artificial food and with small bacteria. The size of the particles ranged from 0.1 to 35 m with special emphasis to the particle diameters between 0.1 and 1 m. In one set of experiments a mixture of differently sized particles was offered as food suspension and the selectivity of filtering was determined by comparing the size spectrum of the particles found in the gut contents with the spectrum in the food suspension. In a second series of experiments suspensions of uniformly sized particles were offered to single animals and their feeding activity was observed directly. In both types of experiments the mesh sizes of the filtering apparatus of the respective animals studied were measured after the experiments by, scanning electron microscopy. The mean sizes of the filter meshes were about 0.4–0.7 m. In all experiments the size of the particles found in the gut or those which caused high feeding activities were larger than the smallest mesh sizes of the filters. As a consequence simple mechanical sieving provides a sufficient explanation for the mechanism of particle retention of the filtering process in Daphnia. D. magna was found to feed with high efficiency on suspended freshwater bacteria, the residual species investigated showed low filtering efficiencies when bacteria were offered as food.The present study was supported by Deutscher Akademischer Austauschdients     

Seasonal patterns of food limitation in Daphnia galeata: separating food quantity and food quality effects

Food limitation for the cladoceran Daphnia galeta was measuredgrowing them with natural lake seston for one growing seasonunder standardized conditions in the laboratory. Growth rateswere related to several measures of food quantity. Particulateorganic carbon (POC) <30 µm was the best predictorof total food availability measured. It was better than chlorophyll-a(Chl-a) of the same size fraction, algal volume calculated fromcell counts or total particle volume from Coulter Counter measurements.Daphnids were also grown with Scenedesmus acutus under the sameconditions and their growth rates were compared. Sestonic carbonconcentrations were related to Scenedesmus carbon concentrations,that provided equal growth rates. The ratio of the carbon concentrationsof both food types (seston, Scenedesmus) represented the foodquality of seston. Thus, the nutritional quality of naturallake seston relative to a standard alga was determined overone growing season. Growth rates of seston fed animals revealedthat they were food limited during long periods of the year.This was also ascertained by standard clutch sizes of fieldanimals. However, the causes of food limitation varied withinthe duration of the studied period. Whereas during the clear-waterphase the food quantity was limiting (while the quality washigh: 100% Scenedesmus), shortly after on 10 June and 17 Junefood conditions were close to threshold concentrations due tolow food quality. For the remainder of the summer, a POC of0.3–0.6 mg C I^–1 with a quality of     

Egg size and clutch size in two Daphnia species grown at different food levels

The effect of three different Scenedesmus food concentrations(0.04, 0.2 and 1 mg C l^–1) on maternal investment wasstudied in two cladoceran species of similar size, Daphnia pulicariaand D.hyalina. It was observed that as food concentration decreased(between 1 and 0.2 mg C l^–1), there was an increase insize, protein content, lipid content, carbon and mass of theegg, while, at the same time, the clutch size of the femalesbecame smaller. Such an increase in ‘per offspring investment’was reflected in an increase in body length, body carbon andbody mass of neonates as the food available for females decreased.However, in D.pulicaria this tendency was not maintained downat 0.04 mg C l^–1 in which there was a decrease of theegg characteristics mentioned above. Although, there are notavailable all the egg and neonate parameters of D.hyalina at0.04 mg C l^–1, the body length of the neonates was largerthan at 0.2 and 1 mg C l^–1. These results show that, asfood diminishes, these two cladoceran species are able to respondby decreasing clutch size, but increasing the size of egg, therebyincreasing the probability of neonate survival. This tendencyis probably maintained until the food concentration is too lowand the females have to reduce the energy allocated for reproduction.     

Egg size and egg mass of Daphnia magna: response to food availability

The influence of different food availability on egg size and egg mass in Daphnia magna Straus was studied in long-term experiments using a flow-through system. Daphnia were either kept a constant high or low food levels or subjected to alternating periods of high food and starvation. Some animals were starved continuously after they had deposited their first clutch of eggs. Eggs were measured and weighed and their density (dry mass per volume) was determined. The results support the model of Glazier (1992), which defines a region of reproductive constraint at very low food concentrations and a region of adaptive response as food concentration increase. Egg sizes were largest under continuously low food concentrations (0.1 mg Cl^–1), which indicates that the maximum of Glazier's non-linear response curve is at very low food levels. Eggs produced during starvation were small, probably as a result of reproductive constraints. Egg density was about 0.37 mg dry weight mm^–1 and did not differ between treatments.     

The heart of Daphnia magna: effects of four cardioactive drugs

We used Daphnia magna bioassays to determine the LC(50) and the effects on the heart of the cardioactive drugs ouabain, verapamil, metaproterenol and metoprolol. Distinctions were made between the pharmacological and toxicological effects of these drugs and the adequacy of physicochemical characteristics of its habitat (reconstituted water). Video microscopy and digital image processing were used to study the pharmacological effects on the heart. D. magna exhibited the expected sensitivity to the reference toxicant sodium dodecyl sulfate with a LC(50) of 15.6+/-4.5 mg/l. All drugs were toxic with 48 h-LC(50) of 2.03 mg/l ouabain, 7.04 mg/l verapamil, 32.45 mg/l metaproterenol and 76.21 mg/l metoprolol. Ouabain was the most toxic and caused a positive concentration-dependent inotropic effect. Verapamil caused positive chronotropic and inotropic effects, while metaproterenol showed positive concentration-dependent chronotropic effects at high concentrations (10(-3) and 10(-4) M). Metoprolol induced a positive chronotropic effect at low concentrations (10(-8), 10(-7), 10(-6) M) and a negative chronotropic effect at high concentration (10(-4) M). Ouabain, metaproterenol and metoprolol in D. magna caused similar effects to those produced in mammals. In contrast, verapamil caused opposite effects. The results suggest the presence of Na(+), K(+)-ATPase receptors to verapamil and of non-specific adrenergic receptors in heart of D. magna.     

Life history responses of Daphnia clones to toxic Microcystis at different food levels

We examined how the amount of food of good quality, Scenedrsmusobtusiusculus, influenced the toxic effects of the cyanobacteriumMicrocystu aeruginosa on Daphnia pulex in a 21 day life tableexperiment. Our results show that all life history traits studied,i.e. longevity, growth and reproduction, of D.pulex were negativelyaffected by toxic Microcystis. The detrimental effects on theselife history traits were less pronounced at increasing foodlevels. In addition, D.pule.x clones differed in their lifehistory responses to toxic Microcystis, suggesting that thepresence of toxic cyanobacteria may modify the outcome amongD.pulex clones.     

Temperature and the effects of elemental food quality on Daphnia

1. We examined the responses of two species of Daphnia to changes in food phosphorus (P) content, with animals reared at three different water temperatures. Specifically, we measured mass‐specific growth rate (MSGR), body P content and respiration rate of Daphnia magna and Daphnia pulex acclimatised to 10, 17.5 and 25 °C and fed food carbon : phosphorus (C : P) ratios of either 150 or 500. 2. The responses of these three physiological variables to temperature–food quality interactions were species‐specific. There was a significant interactive effect of temperature and food quality on D. magna, as the greatest proportional effect of food quality on growth was observed at 10 °C and reductions in body P because of low food P content were relatively greater at 25 °C. These effects may reflect the temperature dependence of mechanisms that reduce elemental constraints associated with food quality in D. magna. By contrast, there were no interactive effects between food quality and temperature on MSGR, body P or mass‐specific respiration of D. pulex. 3. It thus appears that temperature can alter food quality effects on Daphnia but the nature of these alterations depends upon the daphniid species and its thermal adaptability. Significant temperature–food quality interactions will complicate efforts to understand zooplankton nutrition in nature and warrant future consideration.     

Clutch size and body size at first reproduction in Daphnia pulicaria at different levels of food and predation

Field data from seven alpine lakes in Serra da Estrela. Portugal.show that reproduction in Daphnia may be as efficiently controlledby fish predation and copepod predation on eggs in brood cavitiesas it is by food limitation. Body length and clutch size estimatesin Daphnia pulicaria revealed high inter- and intra-populationvariability in maturation size (body size at first reproduction).and in number of eggs per clutch. Daphnia at first maturationin lakes stocked with rainbow trout were half the size of thosefound in fishless lakes (body length of 0.86–0.95 and1.55–1.81 mm. respectively). The mean number of eggs perclutch was reduced to a similar degree by food limitation, predationby fish and copepod predation on eggs in brood cavities, butthe underlying mechanisms of this reduction were different.Food limitation caused smaller clutch sizes in all individuals,so variation remained the same. Fish predation caused the selectiveremoval of individuals with maximum clutches, so variation decreased.Copepod predation caused removal of eggs from brood cavitiesof randomly infested females, so that variation increased, particularlyat a high food level when non-infested females carried largeclutches of eggs.     

Sensitization and habituation of the swimming behavior in ascidian larvae to light

Ascidian larvae of Ciona intestinalis change their photic behavior during the course of development. Newly hatched larvae show no response to a light stimulus at any intensity. At 4 hr after hatching, larvae were induced to start to swimming upon the cessation of illumination, and to stop swimming upon the onset of illumination. At a weaker light intensity (5.0 x 10(-3) J/m (2).s), the larvae showed similar responses to either a single stimulus or repeated stimuli of onset and cessation of light until 10 hr after hatching. At a stronger light intensity (3.2 x 10(-1) J/m(2).s), when the stimulus was repeated, they showed sensitization and habituation of the swimming response. At 3 hr after hatching the larvae failed to show any response to an initial stimulus at any intensity of light, but after several repeated stimuli (sensitization) they showed a swimming response at light intensities above 4.0 x 10(-2) J/m (2).s. At 5 hr and with intensity above 1.0 x 10 (-2) J/m(2).s, the larvae showed photoresponses to the first stimulus, but after several repetitions the larvae failed to stop swimming upon the onset of light (habituation). A repeated series of stimuli at stronger intensities of light caused greater habituation; this habituation was retained for about 1 min. Since the larval central nervous system in Ciona is comprised of only about 100 neurons, learning behavior in ascidian larvae should provide insights for a minimal mechanism of memory in vertebrates.     

Predation by Neomysis mercedis: effects of temperature, Daphnia magna size and prey density on ingestion rate and size selectivity

1. Neomysis mercedis predation rates on Daphnia magna were determined under laboratory conditions. There were generally no consistent differences between the number of Daphnia ingested at 10 and 14°C. 2. At each temperature, the number of prey consumed increased with mysid size and decreased with Daphnia size. 3. For small prey the relationship between ingestion rate and prey density represented a Type II functional response. However, for larger prey there was no significant relationship between density of prey and consumption by mysids. 4. The pattern of size-selective predation by Neomysis was studied to test the optimal foraging hypothesis. For prey populations with mixed size classes, the smallest size of prey was consumed most frequently but intermediate size prey provided the greatest biomass. These observations are contrary to our predictions based on calculations of profitability of different sizes of prey.     

Filtering rate of the cladoceran,Daphnia pulex as a function of body size,light and acclimation

Daphnia pulex were raised under nine light intensities (0, 1.7, 3.5, 7, 14, 28, 55, and 110 ft-c), polarized light (6.6 ft-c), and four wavelength ranges.Light intensity significantly affected the relationship between filtering rate and body size of unacclimated animals. The b values were lowest at 7 and 14 ft-c and they increased above 7 ft-c as light intensity increased. There were significant differences among b values and adjusted means for unacclimated animals. Acclimation to their respective conditions resulted in some significant differences between b values and adjusted means. For acclimated animals there were no significant differences among b values but there were some differences among adjusted means. The filtering rates of unacclimated and acclimated animals were lower at 14 ft-c while 1.7 and 3.5 ft-c were generally stimulatory and the effect was more pronounced in larger animals. Light intensities above 28 ft-c tended to suppress the filtering rate of small unacclimated animals and stimulate filtering rate in larger unacclimated animals. After acclimation, intensities above 7 ft-c did not affect the filtering rate of either small or large animals.The effect of polarized light on filtering rate was inseparable from the effect of light intensity, however, acclimation to polarized light resulted in a significantly higher b value.There were no significant effects of wavelength among b values for unacclimated animals and the adjusted mean for blue wavelengths was significantly higher than for violet wavelengths. There were no significant differences among b values or adjusted means for acclimated animals. Acclimation to their respective wavelengths did result in some significant differences between b values and adjusted means. Except for the effects of blue wavelengths on unacclimated animals and red wavelengths on acclimated animals, the effects of wavelengths are inseparable from the effects of light intensity.This paper is part of a Ph.D. Thesis submitted to the Faculty of Graduate School of The University of Kansas.Supported in part by a NSF Summer Traineeship and a grant from the Research Corporation to St. Olaf College, Northfield, Minnesota 55057.     

Transgenerational effects of poor elemental food quality on Daphnia magna

Environmental effects on parents can strongly affect the phenotype of their offspring, which alters the heritability of traits and the offspring’s responses to the environment. We examined whether P limitation of the aquatic invertebrate, Daphnia magna, alters the responses of its offspring to inadequate P nutrition. Mother Daphnia consuming P-poor algal food produced smaller neonates having lower body P content compared to control (P-rich) mothers. These offspring from P-stressed mothers, when fed P-rich food, grew faster and reproduced on the same schedule as those from P-sufficient mothers. In contrast, offspring from P-stressed mothers, when fed P-poor food, grew more slowly and had delayed reproduction compared to their sisters born to control mothers. There was also weak evidence that daughters from P-stressed mothers are more susceptible to infection by the virulent bacterium, Pasteuria ramosa. Our results show that P stress is not only transferred across generations, but also that its effect on the offspring generation varies depending upon the quality of their own environment. Maternal P nutrition can thus determine the nature of offspring responses to food P content and potentially obfuscates relationships between the performance of offspring and their own nutrition. Given that food quality can be highly variable within and among natural environments, our results demonstrate that maternal effects should be included as an additional dimension into studies of how elemental nutrition affects the physiology, ecology, and evolution of animal consumers.     

Seasonal variations in copepod size: effects of temperature, food abundance, and vertebrate predation

Seasonal dry weights of female and male Cyclops bicuspidatusthomasi, Diaptomus ashlandi and Diaptomus minutus were studiedin southeastern Lake Michigan during 1975–1981. Smallestanimals occurred during summer and early fall, and largest animalsin winter and spring, a pattern reported for other copepods.The range of weights for the species and sexes decreased fromwinter to summer, and converged in summer to a value of approximately2 µg/animal. Surface water temperature and abundancesof young-of-the-year (YOY) fish were inversely correlated withweights of males and females of the three copepod species. Standingstocks of important phytoplankton groups, especially pennatediatoms and flagellates, were positively correlated with copepodweight. Seasonal change of copepod body size appears to be morethan simply a function of temperature; seasonal predation byYOY fish may be a factor influencing sizes of adult copepods.     

Chronic responses of different clones of Daphnia longispina (field and ephippia) to different food levels

Resting eggs are a fundamental reproductive strategy among freshwater cladocerans. Under adverse environmental conditions, whole Daphnia populations can disappear from a lake and a new community will arise from ephippial eggs. Since these new populations are subjected to genetic variation, their responses to environmental stress or contaminants can be different from the “original” population. In the present study, life history responses (reproduction and growth) of Daphnia longispina to different food concentrations was studied. Two Daphnia populations were tested: (a) field clones and (b) ephippial clones. Food (Selenastrum capricornutum) concentration was the stressor tested (absence of food, and low to high food concentrations). The results showed that reproductive responses of D. longispina to the tested food concentrations varied among clones, independently from their origin.     

Phenotypic plasticity and constancy of life-history traits in laboratory clones of Daphnia magna Straus: effects of neonatal length

1. The phenotypic constancy of four laboratory Daphnia magna clones in fitness-related life-history traits, such as age and clutch size at maturity, was studied among consecutive experimental runs in differing food environments.
2. A significant part of the observed clonal and genetic-by-environmental variation in age and clutch size at maturity was explained by experimentally uncontrollable variations in neonatal body length.
3. Despite food availability, neonatal length determined the number of instars invested to maturity and thus maturation age. Clonal differences in neonatal length and thus in maturation instar occurrence across environments explained most of the clonal variability observed in maturation age.
4. Although interclonal differences in clutch size existed, most of the phenotypic plasticity observed for clutch size was mediated by clonal differences in neonatal length.
5. Clonal differences in neonatal length and in the occurrence of maturation instars across environments dramatically affected the body length of instar IM-2 where provisioning of eggs take place. Since clutch size is determined from clutch mass and clutch mass was strongly related to the body length of instar IM-2, clonal differences across environments in body length of instar IM-2 mirrored clonal differences across environments in clutch size.
6. The results reported in the present study show that maternally mediated traits such as neonatal length affect how genotypes respond to different environmental selection regimes (genetic-by-environmental interaction). Future research needs to focus on the effects of neonatal length on the heritability or genetic variation of the reaction norms, since prediction of the response to selection is a key research objective in quantitative genetic studies.