Positive effects of UV radiation on a calanoid copepod in a transparent lake: do competition, predation or food availability play a role?

Zooplankton tolerant to ultraviolet radiation (UVR) could beindirectly affected by UVR through interactions with UV-sensitivespecies in the same ecosystem. In Lake Giles, Pennsylvania,USA, the calanoid copepod Leptodiaptomus minutus is more UVRtolerant than the cohabiting species Daphnia catawba and Cyclopsscutifer. We asked whether L. minutus is affected by UV-inducedmortality of a food competitor (D. catawba) or a predator ofits nauplii (C. scutifer). We conducted two in situ enclosureexperiments with six treatments: L. minutus alone, L . minutus+ Daphnia and L. minutus + Cyclops in the presence and absenceof UVR. There were few differences in survival among treatmentsin Experiment 1, which had enhanced food and a cumulative UVR(320 nm) dose of 9.3 kJ m^–2. In Experiment 2, which hadambient food and a UVR (320 nm) dose of 20.0 kJ m^–2, L.minutus survival and reproduction were higher in the +UVR comparedto –UVR, regardless of competitors or predators. Chlorophylla (Chl a) in Experiment 2 was higher in the +UVR than –UVR.While interactions between zooplankton species of differingUVR tolerances are potentially important, these results insteaddemonstrate that the beneficial UVR effect on L. minutus isindependent of concurrent detrimental UVR effects on competitorsand predators. Further research on the phytoplankton communityis necessary to determine whether UVR alleviates bacterial competition,increases nutrient availability or affects phytoplankton byother mechanisms.     

Influence of Solar Ultraviolet Radiation on Early Life-history Stages of the Bluegill Sunfish, Lepomis macrochirus

We examined the effect of UVR on the success of early life history stages of Lepomis macrochirus in two lakes with contrasting underwater UVR environments. We tested the prediction that when exposed to similar amounts of UVR, bluegill embryos from lakes with high UVR environments have lower mortality than those from lakes with low UVR. We measured embryo survival in the two lakes by performing three experiments in which ambient UVR was manipulated. Embryos responded similarly to UVR exposure regardless of the lake of origin. However, UV-B in UVR transparent lakes is a potentially important factor in the success of early life history stages of the bluegill. A survey of the natural nesting depths in the two lakes revealed that the median depth at which bluegills construct their nests was greater in Lake Giles (high underwater UVR environment) than in Lake Lacawac (low underwater UVR environment).The first author is also the senior author     

Effects of Ultraviolet Radiation on Diel Vertical Migration of Crustacean Zooplankton: An in situ Mesocosm Experiment

The objective of this study was to expand the spatial scale of previous experiments on the effects of ultraviolet radiation (UVR) on diel vertical migration (DVM) by freshwater zooplankton. We conducted an in situ mesocosm experiment in highly UVR transparent Lake Giles, Pennsylvania, in which we imposed two treatments: ambient UVR and UVR-shielded. Mesocosms (3440 L, 0.74 m diameter, 8 m deep) were large enough to include a spatial refuge from UVR and permit relatively large-scale DVM. Daphnia catawba adopted a significantly deeper distribution during the day in the ambient UVR treatment compared to the UVR-shielded treatment, but effects of UVR were absent at night. In contrast, DVM by Leptodiaptomus minutus was unaffected by the UVR treatment. In both treatments, Leptodiaptomus minutus were most abundant at the bottom of the mesocosms during the day and exhibited a more uniform distribution across depths at night. These results suggest that UVR, along with temperature, algal resources, and predators, may affect zooplankton DVM in aquatic ecosystems.     

Mixotrophic algae in three ice-covered lakes of the Pocono Mountains, U.S.A.

1. The occurrence and grazing activity of mixotrophic (phagotrophic) algae in three icecovered freshwater lakes of different trophic status were examined (oligotrophic Lake Giles, mesotrophic Lake Lacawac, eutrophic Lake Waynewood), Microbial population densities were low (4.1–7.2 × 10⁵ bacteria ml^?1 and 1.2–2.4 × 10³ nanoplanktonic protists ml^?1). All three nanoplankton communities were dominated by chloroplast-bearing forms (60–96%). 2. Mixotrophs formed up to 48% of the phototrophic nanoplankton in Lake Lacawac and were responsible for up to ～90% of the observed uptake of bacteria-sized particles. The abundance of mixotrophic algae in Lakes Giles and Waynewood were extremely low (3 and 2% of the phototrophic algae, respectively), and heterotrophs dominated nanoplankton bacterivory. 3. The overall impact of nanoplankton feeding activity on the bacterial assemblage was low under the ice in Lakes Giles and Waynewood. Removal rates of bacteria based on our particle uptake experiments were 1.0 and 4.0% of the bacterial standing stock day^?1 in these lakes, respectively. Removal rates were higher in Lake Lacawac and ranged from 4.9 to 11% of the bacterial standing stock day^?1 on 2 successive sampling days.     

An examination of the potential effects of food limitation on the ultraviolet radiation (UVR) tolerance of Diaptomus minutus

• 1 A series of field and laboratory experiments was performed to test the possible influence of food limitation on UVR tolerance in the calanoid copepod Diaptomusminutus.
• 2 Food‐limited animals incubated at 0.5 m in a clear‐water lake for over three days showed no evidence for increased mortality compared to food‐replete animals when exposed to UVR.
• 3 Animals exposed to artificial UVR in the laboratory showed high mortality. There was no evidence that feeding improved UVR tolerance.
• 4 The results of these experiments show that UVR tolerance is not affected by a short‐term lack of energy or a lack of essential nutrients. However, effects of non‐nutritive dietary components such as carotenoids and mycosporine‐like amino acids cannot be excluded.

     

Assessment of grazing by the freshwater copepod Diaptomus minutus using carotenoid pigments: a caution

In order to assess feeding selectivity in freshwater zooplankton, we conducted feeding trials using Diaptomus minutus isolated from two Wisconsin lakes. Copepods were fed an algal assemblage comprised of an equal biomass of a centric diatom, a cryptomonad and a coccal green alga. The total amounts of photopigments were tracked using high-performance liquid chromatography. The removals of carotenoids and a-type phorbins (chlorophyll a and phaeopigments) from feeding suspensions were compared with their presence in the guts of animals, in fecal pellets, and in the final suspension. Diaptomus minutus generally removed either the diatoms primarily or all three algal cells equally. These removals were not reflected in gut extracts of the animals, however, where alloxanthin (marker of cryptomonads) was always present, but where fucoxanthin and diadinoxanthin (markers of diatoms) were never observed. Pigment disappearance was variable for total a-type phorbins, but frequently >90% for carotenoids, particularly for fucoxanthin and diadinoxanthin. Phaeophytin a was the major a-type phorbin detected after gut passage. Our results indicate that evaluations of zooplankton grazing which assume that algal carotenoids remain detectable throughout gut passage must be made with substantial caution and that differences in pigment processing are likely to occur among zooplankton species. Furthermore, grazing experiments designed to evaluate decreases of specific pigments in feeding suspension, rather than their appearance in animals' guts, may prove a more valuable approach to understanding the feeding selectivity of copepods.      

The susceptibility of the moderate halophile Vibrio costicola to heavy metals

Fifty-eight strains of the moderate halophile Vibrio costicola , including both culture collection strains and freshly isolated strains from solar salterns, were examined for their susceptibility to 10 heavy metal ions by using an agar dilution technique. All strains were sensitive to cadmium, copper, silver, zinc and mercury. This latter ion showed the highest activity even at 0·05 mmol/1 metal concentration. On the other hand, all strains were similarly tolerant to lead, and a great proportion of them were also tolerant to nickel (91%) and chromium (88%). Only 44% and 14% of them showed tolerance to arsenate and cobalt, respectively. The majority of strains (96·4%) were multiply metal-tolerant, with three different metal ion tolerances as the major pattern.     

Evolutionary shifts in copepod acid tolerance in an acid-recovering lake indicated by resurrected resting eggs

We employed zooplankton resting eggs to track population-level shifts in acid tolerance over the last century in a boreal shield lake recovering from acidification. Shifts in mean and variance of ecological tolerances of species that occupy ecosystems recovering from anthropogenic stress are important to consider because of their potential to influence eco-evolutionary dynamics at community and ecosystem levels. In a laboratory experiment, we compared juvenile survival of Leptodiaptomus minutus copepods hatched from resting eggs from three time periods (80- to 100-year- old: pre-industrial, 20- to 50-year- old: lake acidification, and 8- to 10-year- old: recovery of lake-water back to pH ≥ 6.0) under several pH treatments. Mitochondrial DNA was used to confirm species identity and to test for population bottlenecks as a possible mechanism to explain our results. We expected that nauplii hatched from eggs deposited prior to industrialization (lake-water pH ≈ 6.0) and from the period of pH recovery (lake-water pH ≥ 6.0) would have lower mean and more variable survival at acidic pH compared to nauplii hatched from the period of peak lake acidification (lake-water pH ≈ 4.7). Our results, which are likely a combination of both genetic and environmental effects, suggest support for this hypothesis. Nauplii hatched from eggs deposited during the period of acidification in George Lake had reduced variation in pH tolerance compared to the recovery period. This was likely driven by strong selection rather than genetic drift because we found no evidence of a population bottleneck. However, we could not detect differences in the variance of naupliar survival between pre-industrial and acidic time periods. Trends in mean naupliar survival from different time periods matched findings from other field-based studies that detected a relationship between lake acidification history and acid tolerance in L. minutus.     

Mycosporine-like amino acids protect the copepod Boeckella titicacae (Harding) against high levels of solar UVR

Studies were conducted in Lake Titicaca, Bolivia (16°S,69°W, 3810 m a.s.l.) to determine the effects of solar ultravioletradiation (UVR; 280–400 nm) and the protective role ofmycosporine-like amino acids (MAAs) in the calanoid copepodBoeckella titicacae (Harding). Zooplankton samples were takenfrom two different radiation environments: Lake Titicaca anda tank with running lake water, which was covered with layersof neutral-density screen for over a year, receiving ~10% ofincident radiation. Samples collected from the lake had significantlyhigher amounts of MAAs compared with those from the tank. Duringshort-term experiments (6–8 h incubations; six radiationtreatments using sharp cut-off Schott filters), significantlyhigher mortality was observed in B. titicacae from the tankthan from the lake. The higher sensitivity of B. titicacae containinglow amounts of MAAs (compared with specimens collected fromthe lake) was assessed through the determination of biologicalweighting functions (BWFs). A 3-day-long experiment performedwith specimens collected from the lake indicated an increasein mortality as a function of UVR exposure, reaching valuesas high as 70% at the end of day 3 for those individuals thatreceived full radiation compared with <10% for those thatreceived only photosynthetically active radiation (PAR; 400–700nm). Concomitantly, a reduction in the concentration of UV-absorbingcompounds as a function of UVR dose and decreasing wavelengthwas determined. The relatively high UVR dose threshold for mortality(~300 kJ m^–2), as well as a comparison with related species,demonstrated the high resistance of B. titicacae to UVR. Ourresults indicate that this resistance is provided by the presenceof protective compounds, i.e. MAAs, which were efficiently bioaccumulatedthrough the diet.     

INTER- AND INTRASPECIFIC VARIATION OF MOSSES IN TOLERANCE TO COPPER AND ZINC

Bryophytes are often viewed as slowly evolving with little genetic variation within and among populations. A study of heavy-metal tolerance was initiated to test the capacity of bryophytes to undergo genetic differentiation in response to natural selection. Tolerance of Funaria hygrometrica to copper and zinc was greater in populations that originated on soil with high concentrations of these metals. Protonemal growth was more inhibited by the metals than was germination, and copper was more toxic than zinc. Zinc and copper tolerances were correlated, but so were the zinc and copper concentrations of native substrates. The pattern of population differentiation for heavy-metal tolerance in this species is much like that of flowering plants. Five populations of Physcomitrium pyriforme, which does not occur on metal-contaminated soil, were all highly tolerant of zinc but extremely intolerant of copper. This species seems to have an inherent tolerance to the former. Significant variation in tolerance to copper and zinc occurred among populations, but tolerance did not correlate with metal contents in native substrates. This pattern differs from that of flowering plants. Normal populations of species that colonize contaminated sites tended to be more tolerant than populations of species that do not colonize such sites. The extensive population differentiation in Funaria hygrometrica augments the evidence from electrophoretic data that there is genetic variation among populations of mosses and liverworts.     

Ultraviolet radiation and bio-optics in Crater Lake,Oregon

Crater Lake, Oregon, is a mid-latitude caldera lake famous for its depth (594 m) and blue color. Recent underwater spectral measurements of solar radiation (300–800 nm) support earlier observations of unusual transparency and extend these to UV-B wavelengths. New data suggest that penetration of solar UVR into Crater Lake has a significant ecological impact. Evidence includes a correlation between water column chlorophyll-a and stratospheric ozone since 1984, the scarcity of organisms in the upper water column, and apparent UV screening pigments in phytoplankton that vary with depth. The lowest UV-B diffuse attenuation coefficients (K _d,320) were similar to those reported for the clearest natural waters elsewhere, and were lower than estimates for pure water published in 1981. Optical proxies for UVR attenuation were correlated with chlorophyll-a concentration (0–30 m) during typical dry summer months from 1984 to 2002. Using all proxies and measurements of UV transparency, decadal and longer cycles were apparent but no long-term trend since the first optical measurement in 1896.     

Chloroxylenol- and triclosan-tolerant bacteria from industrial sources

Potential development of bacterial tolerance to biocides in the industrial environment is examined in this study. Bacteria tolerant to the phenolic-type agent para-chloro-meta-xylenol (PCMX) and the bis-phenol 2,4,4′-trichloro-2′-hydroxydiphenylether (triclosan) were isolated from industrial sources and identified. Minimum inhibitory concentrations (MICs) were determined and compared with those of culture collection (standard) strains. Of around 100 isolates originally obtained, most were naturally tolerant species such as Pseudomonas spp., or showed low tolerance levels. PCMX-tolerant isolates of Pseudomonas stutzeri and triclosan-tolerant isolates of Citrobacter freundii and Acinetobacter johnsonii were retained for further study. Of these, only P. stutzeri and A. johnsonii showed elevated tolerance compared with the standard strains. There was no evidence of tolerance to the other biocide except for Pseudomonas aeruginosa (an intrinsically tolerant microorganism), and tolerances were stable in the absence of selective pressure except for A. johnsonii. Attempts to select or generate increased tolerance in the standard strains were unsuccessful. High tolerances in terms of MIC were not reflected in terms of lethal effects. This study did not produce any evidence suggesting that the presence of residual biocide concentrations in the industrial environment promotes the emergence of bacterial tolerance for them. Received 01 February 2002/ Accepted in revised form 09 August 2002     

Effects of nutrients and dissolved organic matter on the response of phytoplankton to ultraviolet radiation: experimental comparison in spring versus summer

The effects of nutrients and dissolved organic matter (DOM) on the response of phytoplankton community structure to ultraviolet radiation (UVR) was studied using natural phytoplankton assemblages from Lake Giles (Northeastern Pennsylvania), a temperate, oligotrophic, highly UVR-transparent lake. Microcosm experiments were conducted in 1-l bags in the spring and summer. A factorial design was used, with two UVR treatments (ambient and reduced), two nutrient treatments (control with no nutrients added, and nitrogen and phosphorus addition together), and two DOM treatments (control of 1 mg l⁻¹ and doubled). In April, UVR affected the overall phytoplankton community structure, causing a shift in the dominant species. Significant interactive effects of UVR × nutrients and UVR × DOM were found on total phytoplankton biovolumes. In July, all taxa responded positively to the N + P addition, and were affected differentially by the UVR treatments. The initial communities varied in April and July, but Synura sp. and Chroomonas sp. were present in both seasons. Synura sp. responded positively to the addition of DOM in April and the reduction of UVR in July. Chroomonas sp. responded positively to the reduction of UVR in April and the addition of nutrients in July. The differential sensitivity of these two species suggests that changing environmental factors between spring and summer promoted differences in the relative importance of UVR in changing phytoplankton community structure. Handling editor: Luigi Naselli-Flores     

Inhibition of phytoplankton photosynthesis by solar ultraviolet radiation: studies in Lake Titicaca, Bolivia

1. Lake Titicaca is a large, high altitude (3810 m a.s.l.) tropical lake (16°S, 68°W) that lies on the border of Bolivia and Perú, receiving high fluxes of ultraviolet radiation (UVR) throughout the year. Our studies were conducted during September of 1997 with the main objective of studying the impact of solar UVR upon phytoplankton photosynthesis.
2. Water samples were taken daily and incubated in situ (down to 14 m depth) under three radiation treatments to study the relative responses to PAR (Photosynthetic Available Radiation, 400–700 nm), UV-A (320–400 nm), and UV-B (280–320 nm) radiation.
3. Photosynthetic inhibition by UVR in surface waters was about 80%, with UV-A accounting for 60% and UV-B for 20%; the inhibition by high levels of PAR was less than 20%. The inhibition due to UVR decreased with depth so that there were no significant differences between treatments at 8.5 m depth.
4. The amount of inhibition per unit energy received by phytoplankton indicates that even though there was a significant inhibition of photosynthesis due to UVR, species in Lake Titicaca seem to be better adapted than species in high latitude environments.
5. The cellular concentration of UV-absorbing compounds, a possible mechanism of photoadaptation, was low in phytoplanktonic species. However, they were abundant in zooplankton, suggesting a high rate of bioaccumulation through the diet.     

Divergence in DNA photorepair efficiency among genotypes from contrasting UV radiation environments in nature

Populations of organisms routinely face abiotic selection pressures, and a central goal of evolutionary biology is to understand the mechanistic underpinnings of adaptive phenotypes. Ultraviolet radiation (UVR) is one of earth's most pervasive environmental stressors, potentially damaging DNA in any organism exposed to solar radiation. We explored mechanisms underlying differential survival following UVR exposure in genotypes of the water flea Daphnia melanica derived from natural ponds of differing UVR intensity. The UVR tolerance of a D. melanica genotype from a high‐UVR habitat depended on the presence of visible and UV‐A light wavelengths necessary for photoenzymatic repair of DNA damage, a repair pathway widely shared across the tree of life. We then measured the acquisition and repair of cyclobutane pyrimidine dimers, the primary form of UVR‐caused DNA damage, in D. melanica DNA following experimental UVR exposure. We demonstrate that genotypes from high‐UVR habitats repair DNA damage faster than genotypes from low‐UVR habitats in the presence of visible and UV‐A radiation necessary for photoenzymatic repair, but not in dark treatments. Because differences in repair rate only occurred in the presence of visible and UV‐A radiation, we conclude that differing rates of DNA repair, and therefore differential UVR tolerance, are a consequence of variation in photoenzymatic repair efficiency. We then rule out a simple gene expression hypothesis for the molecular basis of differing repair efficiency, as expression of the CPD photolyase gene photorepair did not differ among D. melanica lineages, in both the presence and absence of UVR.     

Cold and heat tolerance of drosophilid flies with reference to their latitudinal distributions

The relation between thermal tolerance and latitudinal distribution was studied with 30 drosophilid species collected from the cool-temperate region (Sapporo), the warm-temperate region (Tokyo and Kyoto) and the subtropical region (Iriomote island) in Japan. In addition, intraspecific variation was examined for five species collected from two localities. The subtropical strains of Scaptodrosophila coracina, Drosophila bizonata and D. daruma were less tolerant to cold than their temperate strains. However, the difference of cold tolerance between these two geographic strains was much smaller than the difference between the species restricted to the subtropical region and those occurring in the temperate region. In D. auraria and D. suzukii, no difference was observed in thermal tolerance between their cool- and warm-temperate strains. Thus, geographic variation in thermal tolerance within species was low or negligible. Interspecific comparisons by phylogenetic independent contrasts revealed that species which had the northern boundaries of their distributions at higher latitudes were generally more tolerant to cold than those which had their boundaries at lower latitudes. However, the data for some species did not agree with this trend. The use of man-protected warm places for overwintering, competition or predation would also affect their distributions. It also appeared that species which had their southern boundaries at higher latitudes were generally more cold-tolerant. The acquisition of cold tolerance may lower a flys capacity to compete, survive or reproduce in warmer climates. On the other hand, no relation was observed between heat tolerance and latitudinal distribution. Heat tolerance was higher in species inhabiting openlands or the forest canopy than in those inhabiting the forest understorey.     

Comparison of Plankton-Water Chemistry Relationships in Three Acid Stressed Lakes in Nova Scotia,Canada

A field program was conducted in 1980 and 1981 on three lakes in Kejimkujik National Park, Nova Scotia, to study plankton-water chemistry relationships in a region of potential acid precipitation stress. Among lake and between year comparisons were made. The three study sites represent two different types of lakes. Kejimkujik Lake (pH 4.8) and Pebbleloggitch Lake (pH 4.5) are both high-colored, dystrophic acid lakes. Beaverskin Lake is a clear, oligotrophic lake which is not as acidic (pH 5.4). The phytoplankton of Kejimkujik Lake was dominated by diatoms, while Pebbleloggitch Lake was dominated by chlorophytes and chrysophytes, and Beaverskin Lake was dominated by cyanophytes. Kejimkujik Lake had the highest total algal cell volume per liter, and Pebbleloggitch Lake the lowest. Rotifer populations composed the majority of the zooplankton communities, while the crustacean zooplankton were dominated by the acid-tolerant copepod Diaptomus minutus. Kejimkujik Lake had the lowest zooplankton biomass and Pebbleloggitch Lake the highest zooplankton biomass per m³. A pattern of highest algal density in Beaverskin Lake, but highest zooplankton density in Pebbleloggitch Lake emerged in both years. This is likely an effect of abundant, but unsuitable food (blue-green algae) in Beaverskin Lake and important detrital food resources in Pebbleloggitch Lake in both years. Multiple regression analysis of water chemistry variables with plankton species produced many significant effects, but failed to show clear patterns. Cause and effect relationships in aquatic ecosystems are poorly delineated by such techniques.     

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.     

Life histories of the copepods Pseudocalanus minutus, P. acuspes (Calanoida) and Oithona similis (Cyclopoida) in the Arctic Kongsfjorden (Svalbard)

The year-round variation in abundance and stage-specific (vertical) distribution of Pseudocalanus minutus and Oithona similis was studied in the Arctic Kongsfjorden, Svalbard. Maxima of vertically integrated abundance were found in November with 111,297 ind m⁻² for P. minutus and 704,633 ind m⁻² for O. similis. Minimum abundances comprised 1,088 ind m⁻² and 4,483 ind m⁻² in June for P. minutus and O. similis, respectively. The congener P. acuspes only occurred in low numbers (15–213 ind m⁻²), and successful reproduction was debatable. Reproduction of P. minutus took place in May/June, and stage distribution revealed a 1-year life cycle with copepodids CIII, CIV, and CV as the overwintering stages. Oithona similis exhibited two main reproductive peaks in June and August/September, respectively. Moreover, it reproduced more or less continuously throughout the whole year with all stages occurring during the entire sampling period, suggesting two generations per year. Both species migrated towards greater depth in November, but O. similis preferred to stay longer in the upper 100 m as compared to Pseudocalanus. The reproduction of the two species in Kongsfjorden seemed to be linked to phytoplankton dynamics.