Patterns of CO2 concentration and inorganic carbon limitation of phytoplankton biomass in agriculturally eutrophic lakes.

Lake eutrophication is a pervasive problem globally, particularly serious in agricultural and densely populated areas. Whenever nutrients nitrogen and phosphorus do not limit phytoplankton growth directly, high growth rates will rapidly lead to biomass increases causing self-shading and light-limitation, and eventually CO2 depletion. The paradigm of phytoplankton limitation by nutrients and light is so pervasively established, that the lack of nutrient limitation is ordinarily interpreted as sufficient evidence for the condition of light limitation, without considering the possibility of limitation by inorganic carbon. Here, we firstly evaluated how frequently CO2 undersaturation occurs in a set of eutrophic lakes in the Pampa plains. Our results confirm that conditions of CO2 undersaturation develop much more frequently (yearly 34%, summer 44%) in these agriculturally impacted lakes than in deep, temperate lakes in forested watersheds. Secondly, we used Generalized Additive Models to fit trends in CO2 concentration considering three drivers: total incident irradiance, chlorophyll a concentration, and lake depth; in eight multi-year datasets from eutrophic lakes from Europe, North and South America, Asia and New Zealand. CO2 depletion was more often observed at high irradiance levels, and shallow water. CO2 depletion also occurred at high chlorophyll concentration. Finally, we identified occurrences of light- and carbon-limitation at the whole-lake scale. The different responses of chlorophyll a and CO2 allowed us to develop criteria for detecting conditions of CO2 limitation. For the first time, we provided whole-lake evidence of carbon limitation of phytoplankton biomass. CO2 increases and eutrophication represent two major and converging environmental problems that have additive and contrasting effects, promoting phytoplankton, and also leading to carbon depletion. Their interactions deserve further exploration and imaginative approaches to deal with their effects.

Field evidence supports former experimental claims on the stimulatory effect of glyphosate on picocyanobacteria communities.

Glyphosate-based herbicides are the most commonly used herbicide worldwide. Although glyphosate is known to be toxic to aquatic organisms, it can also have stimulatory effects on small-size (ø <2 µm) cyanobacteria (Pcy) able to metabolize and degrade glyphosate and AMPA. Several previous experimental studies in micro- and mesocosms reported increases of Pcy abundance in response to glyphosate additions, but comparable field evidence is presently unavailable. We surveyed a large geographical area in order to collect information on Pcy abundance from lakes within the Pampa region (with over three decades of glyphosate usage) and lakes from Patagonia (with virtually no history of glyphosate usage). Fifty-two Pampean lakes and 24 Patagonian lakes were surveyed. We used three indicators of glyphosate impact: herbicide concentration, the presence of phosphonate metabolism genes (responsible for glyphosate and AMPA degradation) in environmental DNA samples, and descriptors of land use in the surrounding area of each lake. We addressed three questions: (1) is there field evidence of stimulatory effects of glyphosate on picocyanobacteria abundance? (2) is the magnitude of the effects of glyphosate in natural systems comparable to that reported under controlled experimental conditions? and (3), how do the effects of glyphosate compare to the effects of other potential environmental drivers of Pcy biomass? The collected evidence is consistent with the hypothesis that long-term agricultural practices relying on glyphosate-based technologies had important effects on freshwater microbial communities, particularly by promoting increases in picocyanobacteria abundance.

Rotifer dynamics in three shallow lakes from the Salado river watershed (Argentina): the potential modulating role of incident solar radiation.

In turbid Pampean lakes, incident solar radiation is a major driver of plankton seasonal dynamics. Higher light availability in summer translates into higher primary production, and therefore more food for zooplankton grazers. However, experimental evidence suggests that food produced under the high irradiance conditions prevailing in summer are less suitable to sustain rotifer population growth than that produced under the lower irradiance conditions typical of winter. Here, we analysed time series datasets corresponding to three shallow lakes from the Salado river watershed. This analysis provided evidence for similar seasonal patterns of rotifer relative abundance over a large geographic area. In addition, we performed life table experiments to test the hypothesis that natural seston produced in winter could sustain higher population growth rates than seston produced in summer. We suggest that the natural seasonal changes in temperature and food generate successive time windows, which may be capitalized by the different grazer species, resulting in predictable phenology of grazer populations.

Contrasting patterns of MAAs accumulation in two populations of the copepod Boeckella gracilipes.

The bio-accumulation of mycosporine-like amino acids (MAAs) is common in planktonic copepods that inhabit environments exposed to high levels of solar radiation. MAAs accumulation in copepods can be affected both by extrinsic (environmental) and intrinsic factors (local adaptation, genotype, etc.). Laboratory experiments were performed to study the bio-accumulation of MAAs in two geographically-isolated populations of Boeckella gracilipes from a mountain and a piedmont lake of North Patagonia. We performed two series of 10-day incubations of B. gracilipes from the different lakes applying two radiation conditions (PAR + UVR and darkness), at five different temperatures (5 to 20 °C) and providing a MAA-free flagellate as food. We assumed that differences in final MAAs concentrations between copepod populations should be exclusively due to environmental factors, and that any difference in the patterns of MAAs accumulation should exclusively arise from differences in MAAs concentration at the time of collection. MAAs concentration was three fold higher in B. gracilipes from Lake Verde than in copepods from the Lake Morenito. The MAAs suite was dominated (∼90%) by a combination of porphyra-334 and mycosporine-glycine in copepods from Lake Verde, and porphyra-334 and MAA-332 in those from Lake Morenito. Two exclusive MAA compounds were identified, mycosporine-glycine in copepods from Lake Verde and shinorine in the copepod population from Lake Morenito. Laboratory experiments showed that: (i) exposure to PAR + UVR stimulated the accumulation of MAAs in both copepod populations; (ii) temperature affected the response of MAAs and, remarkably, low temperatures stimulated MAAs accumulation even in dark incubations, (iii) the response to radiation and temperature in MAAs accumulation was more pronounced in the population with low initial MAAs than in the population with high initial MAAs concentrations. The differences in intrinsic factors between B. gracilipes populations, such as local adaptation to contrasting UV and temperature scenarios, among others, appear to play an important role in determining levels and patterns of MAAs accumulation in B. gracilipes.

Mycosporine-like amino acids in freshwater copepods: potential sources and some factors that affect their bioaccumulation.

Mycosporine-like amino acids (MAAs) are ubiquitous photoprotective compounds in aquatic environments. MAAs are synthesized by a wide variety of organisms (i.e. bacteria, fungi and algae) and their production is photoinducible by ultraviolet radiation (UVR) (280-400 nm) and/or photosynthetically active radiation (400-750 nm). Most animals however, are unable to synthesize MAAs and must acquire these compounds through their diet or from symbiotic organisms. In this paper, we investigate the possible sources of MAAs and factors (temperature and initial MAA concentration) that may affect their bioaccumulation in freshwater copepods. We found that MAA accumulation may occur even if the copepods are cultured on a MAA-free diet. In addition, we found that the bacteriostatic antibiotic, chloramphenicol, inhibits the bioaccumulation of MAAs. These two pieces of evidence suggest that the source of MAAs in these copepods may be prokaryotic organisms in close association with the animals. The two factors investigated in this study, temperature and initial MAA concentrations, were found to affect the rates at which MAAs are accumulated. Temperature had positive effects on both uptake and elimination rates. On the other hand, the rate of uptake decreased at the highest assayed initial MAA concentration, probably because the concentration of MAAs was already close to saturation.

Quality of UVR exposure for different biological systems along a latitudinal gradient.

The exposure of organisms to ultraviolet radiation (UVR) is characterized by the climatology (annual cycle) and the variance (anomalies) of biologically-weighted irradiances at eight geographical locations in austral South America, from 1995-2002. The net effect of UVR on biological systems is a result of the balance of damage and repair which depends on intensity and duration of irradiance and is modulated by its variability. The emphasis in this study is on day-to-day variability, a time scale of importance to adaptive strategies that counteract UVR damage. The irradiances were weighted with DNA- and phytoplankton photosynthesis-action spectra. Low latitude sites show high average UVR. For all sites, the frequency of days with above average irradiances is higher than below average irradiances. Persistence in anomalies is generally low (

UV radiation as a potential driving force for zooplankton community structure in Patagonian lakes.

This article explores the potential role of UV radiation (UVR) as an influence on zooplankton communities. In the first section we provide a general overview of UVR effects on freshwater zooplankton, with an emphasis on Argentine and Chilean environments. In the second section we present the results of a survey involving 53 temperate lakes across a gradient of UVR exposure to determine patterns of species richness and specific diversity. These community characteristics decreased at high potential UVR exposure (i.e. high mean water column irradiance or low lake optical density). A threshold value of mean water column irradiance of approximately 10% of the surface level seems to limit both richness and diversity to minimum values. On the basis of the collected evidence it is not possible to definitely conclude that UVR rather than another covarying factor is responsible for the decrease in specific diversity observed at the lowest end of lake optical depth. However, lakes with values above the previous threshold are likely to exhibit highly depauperate zooplankton communities regardless of the mechanism. As a cautionary note we suggest that changes in the optical characteristics (i.e. changes due to atmospheric conditions, precipitation patterns or vertical displacement of the tree line) may result in sudden shifts in zooplankton community structure.

Mycosporines from freshwater yeasts: a trophic cul-de-sac?

Mycosporine-like amino-acids (MAAs) are found in aquatic bacteria, algae, and animals. A related compound, the mycosporine-glutaminol-glucoside (myc-glu-glu), has recently been reported in freshwater yeasts. Although animals depend on other organisms as their source of MAAs, they can efficiently accumulate them in their tissues. In this work we assessed the potential transfer of the yeast mycosporine myc-glu-glu from the diet into the copepod Boeckella antiqua and the ciliate Paramecium bursaria. For this purpose, we performed experiments to study the feeding of B. antiqua and P. bursaria on the yeast Rhodotorula minuta and their ability to bioaccumulate myc-glu-glu. Bioaccumulation of myc-glu-glu in B. antiqua was assessed through long-term factorial experiments manipulating the diet (Chlamydomonas reinhardii and C. reinhardii + yeasts) and radiation exposure (PAR and PAR + UVR). Shorter term experiments were designed in the case of P. bursaria. The composition and concentration of MAAs in the diet and in the consumers were determined by HPLC analyses. Our results showed that even though both consumers ingested yeast cells, they were unable to accumulate myc-glu-glu. Moreover, when exposed to conditions that stimulated the accumulation of photoprotective compounds (i.e. UVR exposure), an increase in MAAs concentration occurred in copepods fed C. reinhardii plus yeasts as well as in those fed only C. reinhardii. This suggests that the copepods were able to modify their tissue concentrations of MAAs in response to environmental clues but also that the contribution of yeast mycosporines to total MAAs concentration was negligible.

Occurrence of photoprotective compounds in yeasts from freshwater ecosystems of northwestern Patagonia (Argentina).

In this paper we present the results of research on the occurrence, induction and role of photoprotective compounds (PPCs) present in native aquatic yeasts from freshwater Patagonian ecosystems. We focus on the effect of UV radiation (UVR) as a factor that controls the level of photoprotection of yeasts, and explore its potential significance in shaping yeast distributional patterns. The research presented here combines field surveys and laboratory work, including the isolation and culture of native yeasts strains, and laboratory assays under different radiation conditions. The results obtained suggest that yeasts are common dwellers of oligotrophic Patagonian water bodies, and provide the first evidence of the distribution of PPC (carotenoid and mycosporine)-producing yeasts in temperate freshwaters. A greater proportion of carotenogenic yeasts were observed in high-elevation lakes. The yeast strains isolated from these environments were found to produce higher amounts of mycosporines (MYCs), and to present higher tolerance to UVB exposure than those from piedmont lakes. Patagonian yeasts have only one type of MYC, mycosporine-glutaminol-glucoside (myc-glu-glu), which seems common to all other yeasts. By analyzing the production of myc-glu-glu in a large number of yeasts belonging to different taxonomic groups, we propose that this compound may have potential use as a chemotaxonomic marker in yeast systematics. Collectively, our work reveals that in Patagonian freshwater yeasts there is an apparent relationship between the ability to produce PPCs, their tolerance to UV exposure and their success in colonizing habitats highly exposed to UVR.

Mycosporine-like amino acids in planktonic organisms living under different UV exposure conditions in Patagonian lakes.

Mycosporine-like amino acids (MAAs) were studied in zooplankton from 13 Argentinian lakes covering a broad range in altitude, maximum depth and physico-chemical properties of the water. Four to nine different MAAs (predominantly porphyra-334 and shinorine) were found in the copepods Boeckella gibbosa, B. gracilipes, B. meteoris and Parabroteas sarsi, and in the ciliate Stentor amethystinus, while MAAs were undetectable in the cladoceran Daphnia middendorffiana. Among the different copepods, maximum MAA concentrations accounted for 0.25-1.31% of the dry weight, and contents were generally about three to seven times (up to 43 times) higher in the animals living in the clearest lakes compared to those occurring in low-UV systems. This variability in the content of MAAs was related to the lake altitude (r(2) = 0.71), and the fraction of the water column to which 1% of the surface UV radiation at 320 nm penetrated (r(2) = 0.57). Our data therefore underscore the role of MAAs as sunscreens to decrease the potential negative effects of solar radiation, but they also indicate that other environmental factors besides UV transparency play a role in determining MAA concentrations. One lake was selected to obtain additional information on the qualitative composition of MAAs in seston of <100 µm between two sampling sites and over a 2 month study period (austral summer). Six different MAAs were detected in the samples, with porphyra-334 and palythine being predominant. In the copepods collected simultaneously, there was low variation in MAA concentrations between the two sites and over time. Thus, our results suggest that under similar UV exposure conditions MAA contents of planktonic organisms show low temporal variation.

Assessing sublethal effects of ultraviolet radiation in juvenile rainbow trout (Oncorhynchus mykiss).

This paper presents a method for assessing sublethal effects of ultraviolet radiation in fish. The method is based on the assumption that UV-B exposure is stressful to fish. We measured oxygen consumption in UV-B-irradiated rainbow trout juveniles using a UV-transparent respirometer chamber. In addition, we simultaneously monitored fish behavior by videotaping. We found a direct relationship between UV-B irradiance and the percentage increase in oxygen consumption. Increased swimming activity and restless behavior were also noted under UV-B irradiation. The actual mechanisms by which fish perceive UV radiation and a behavioral reaction is triggered are unknown. Based on the present study, a response to UV-A irradiation cannot be ruled out.

Replicated mesocosm study on the role of natural ultraviolet radiation in high CDOM, shallow lakes.

The role of ultraviolet radiation on shallow, high CDOM (colored dissolved organic matter) lakes was investigated during two consecutive summers (1999 and 2000) in replicated mesocosms (rectangular fiberglass tanks). Each tank (volume: 300 L; depth: 40 cm) was covered with a layer (approximately 3 cm) of sediment from lake El Toro (40 degrees 14' S; 70 degrees 22' W) and filled with filtered water. The experimental design consisted of two treatments: full natural radiation (UV-exposed) and natural radiation without ultraviolet radiation (UV-shielded). UV-exposed and UV-shielded treatments differed in most studied variables as revealed by repeated measures ANOVA. UV-exposed tanks displayed lower CDOM levels (dissolved absorbance) of lower average molecular size (absorbance ratio between 250 and 365 nm), higher bacterial biomass, and lower chlorophyll a concentration. The effect on consumers (rotifers and crustaceans) was less noticeable. The results are consistent with UV stimulation of bacteria production mediated by higher rates of CDOM photobleaching, and the photoinhibition of planktonic algae. Thus, a major effect of UVR in shallow, high CDOM ecosystems appears to be the stimulation of heterotrophic pathways and a simultaneous inhibition of photoautotrophs.

A field and laboratory study on factors affecting polymorphism in the rotifer Keratella tropica.

We present data on polymorphism of the rotifer Keratella tropica from an outdoor, mesoscale experiment and a series of laboratory induction assays. In the outdoor experiment the biomass of planktonic crustaceans was noticeably depressed by larval fish predation, and associated with this depression K. tropica underwent a striking reduction of caudal spines. Subsequent laboratory studies showed that the positive association between crustacean biomass and caudal spine length was a cause and effect relationship. This is the first record of rotifer morphological change as an indirect effect of fish predation. In laboratory experiments filtrates of monospecific cultures of a cyclopoid copepod, a calanoid copepod and 5 cladocerans induced a remarkable spine development. Morphological induction showed a direct relationship with the concentration of crustaceans, both under field and laboratory conditions. Long spines were found to be strong deterrents against small predators (Acanthocyclops robustus copepodites), but were useless against large ones (females of the same species). The morphotypes of K. tropica obtained by experimental induction from a single clone encompass much of the worldwide variation of the species.