Abstract. Global change, together with human activities, has resulted in increasing amounts of organic material (including nutrients) that water bodies receive. This input further attenuates the penetration of solar radiation, leading to the view that opaque lakes are more "protected" from solar ultraviolet radiation (UVR) than clear ones. Vertical mixing, however, complicates this view as cells are exposed to fluctuating radiation regimes, for which the effects have, in general, been neglected. Furthermore, the combined impacts of mixing, together with those of UVR and nutrient inputs are virtually unknown. In this study, we carried out complex in situ experiments in three high mountain lakes of Spain (Lake Enol in the National Park Picos de Europa, Asturias, and lakes Las Yeguas and La Caldera in the National Park Sierra Nevada, Granada), used as model ecosystems to evaluate the joint impact of these climate change variables. The main goal of this study was to address the question of how short-term pulses of nutrient inputs, together with vertical mixing and increased UVR fluxes modify the photosynthetic responses of phytoplankton. The experimentation consisted in all possible combinations of the following treatments: (a) solar radiation: UVR + PAR (280–700 nm) versus PAR (photosynthetically active radiation) alone (400–700 nm); (b) nutrient addition (phosphorus (P) and nitrogen (N)): ambient versus addition (P to reach to a final concentration of 30 μg P L−1, and N to reach N:P molar ratio of 31); and (c) mixing: mixed (one rotation from surface to 3 m depth (speed of 1 m 4 min−1, total of 10 cycles)) versus static. Our findings suggest that under ambient nutrient conditions there is a synergistic effect between vertical mixing and UVR, increasing phytoplankton photosynthetic inhibition and excretion of organic carbon (EOC) from opaque lakes as compared to algae that received constant mean irradiance within the epilimnion. The opposite occurs in clear lakes where antagonistic effects were determined, with mixing partially counteracting the negative effects of UVR. Nutrient input, mimicking atmospheric pulses from Saharan dust, reversed this effect and clear lakes became more inhibited during mixing, while opaque lakes benefited from the fluctuating irradiance regime. These climate change related scenarios of nutrient input and increased mixing, would not only affect photosynthesis and production in lakes, but might also further influence the microbial loop and trophic interactions via enhanced EOC under fluctuating UVR exposure.

Some of the most important effects of global change on coastal marine systems include increasing nutrient inputs and higher levels of ultraviolet radiation (UVR, 280–400 nm), which could affect primary producers, a key trophic link to the functioning of marine food webs. However, interactive effects of both factors on the phytoplankton community have not been assessed for the Mediterranean Sea. An in situ factorial experiment, with two levels of ultraviolet solar radiation (UVR+PAR vs. PAR) and nutrients (control vs. P-enriched), was performed to evaluate single and UVR×P effects on metabolic, enzymatic, stoichiometric and structural phytoplanktonic variables. While most phytoplankton variables were not affected by UVR, dissolved phosphatase (APAEX) and algal P content increased in the presence of UVR, which was interpreted as an acclimation mechanism of algae to oligotrophic marine waters. Synergistic UVR×P interactive effects were positive on photosynthetic variables (i.e., maximal electron transport rate, ETRmax), but negative on primary production and phytoplankton biomass because the pulse of P unmasked the inhibitory effect of UVR. This unmasking effect might be related to greater photodamage caused by an excess of electron flux after a P pulse (higher ETRmax) without an efficient release of carbon as the mechanism to dissipate the reducing power of photosynthetic electron transport.

From an extensive study, we determined that heterotrophic bacterial production (HBP) variance in Sierra Nevada (Spain) lakes was explained mainly by excretion of organic carbon by algae (EOC), underlining a bacterial dependence on algal carbon. Subsequently, we studied how the interaction among global change factors such as ultraviolet radiation (UVR), nutrient inputs, and increased temperature affected this phytoplankton-bacteria coupling through in situ factorial experiments in two model high-mountain lakes, La Caldera, and Las Yeguas. Bacterioplankton were more sensitive than phytoplankton because the joint action of increased temperature and nutrient-addition unmasked an inhibitory UVR effect on HBP while reducing the inhibitory UVR effect on primary production (PP) (in La Caldera) or augmenting the net PP values (in Las Yeguas). The interaction among the three factors had a different effect on phytoplankton-bacteria coupling depending on the lake. Thus, in the colder lake (La Caldera), EOC was not adequate to meet the bacterial carbon demand (BCD), leading to a mismatch in phytoplankton-bacteria coupling. Contrarily, in the warmer lake (Las Yeguas), the phytoplankton-bacteria coupling was accentuated by the interaction among the three factors, with EOC exceeding BCD. These contrasting responses of phytoplankton-bacteria coupling may affect the microbial loop development, becoming reinforced in warmer and less UVR-transparent highmountain lakes, but weakened in colder and more UVR-transparent high-mountain lakes, with implications in the C-flux of these sentinel ecosystems in a scenario of global change.

Abstract. An indirect effect of global warming is a reduction in the depth of the upper mixed layer (UML) causing organisms to be exposed to higher levels of ultraviolet (UVR, 280-400 nm) and photosynthetically active radiation (PAR, 400-700 nm). This can affect primary and bacterial production as well as the commensalistic phytoplankton-bacteria relationship. The combined effects of UVR and reduction in the depth of the UML were assessed on variables related to the metabolism of phytoplankton and bacteria, during in situ experiments performed with natural pico-and nanoplankton communities from two oligotrophic lakes with contrasting UVR transparency (high-UVR versus low-UVR waters) of southern Spain. The negative UVR effects on epilimnetic primary production (PP) and on heterotrophic bacterial production (HBP), intensified under increased stratification, were higher in the low-UVR than in the high-UVR lake, and stronger on the phytoplanktonic than on the heterotrophic bacterial communities. Under UVR and increased stratification, the commensalistic phytoplankton-bacteria relationship was strengthened in the high-UVR lake where excretion of organic carbon (EOC) rates exceeded the bacterial carbon demand (BCD; i.e., BCD : EOC(%) ratio < 100). This did not occur in the low-UVR lake (i.e., BCD : EOC(%) ratio > 100). The greater UVR damage to phytoplankton and bacteria and the weakening of their commensalistic interaction found in the low-UVR lake indicates that these ecosystems would be especially vulnerable to UVR and increased stratification as stressors related to global climate change. Thus, our findings may have important implications for the carbon cycle in oligotrophic lakes of the Mediterranean region.

Nutrient inputs and ultraviolet radiation (UVR) are global factors affecting the structure and functioning of aquatic ecosystems, particularly clear-water ecosystems. We performed experiments in two model lakes highly exposed to UVR fluxes in order to test the effect that future increases in mineral nutrients transported by dust aerosol might exert on primary producers depending on the likelihood of atmospheric inputs. Lake La Caldera (Northern Hemisphere) has been receiving recurrent dust inputs from the Sahara Desert while lake Los Cántaros (Southern Hemisphere) has been less affected by dust aerosol. UVR × Nutrient synergistically stimulated primary production (PP), chlorophyll a (Chl a), with a smaller increase in phytoplanktonic biomass in La Caldera, but not in Los Cántaros, where nutrient addition unmasked the UVR inhibitory effect on phytoplankton. The proportional decrease of mixotrophic nanoflagellates (MNFs) after the nutrient pulse (in Los Cántaros) and the long-term decline of MNFs in La Caldera associated with the increase in aerosol-dust intrusions from the Sahara during the last 40 years suggest that a future scenario of intensified aerosol events from desert and desertified areas would not only reduce functional diversity with the decline of MNFs, but would ultimately alter the C flux towards the grazing chain in oligotrophic ecosystems.

Abstract. An indirect effect of global warming is the shallowing epilimnion, causing organisms to be exposed to higher levels of ultraviolet (UVR, 280–400 nm) and photosynthetically active radiation (PAR, 400–700 nm), which could affect primary and bacterial production as well as the commensalistic algal-bacterial relationship. The combined effects of UVR and reduction in the depth of the upper mixed layer (UML) were assessed on variables related to the metabolism of algae and bacteria, during in situ experiments performed with natural microplanktonic communities from two oligotrophic lakes with contrasting UVR-transparency (clear vs. opaque) of southern Spain. The negative UVR effects on epilimnetic primary production (PP) and on heterotrophic bacterial production (HBP), intensified by high mean irradiances, were higher in the UVR-opaque than in the UVR-clear lake, and stronger on the algae than on the heterotrophic bacterial communities. Under UVR and high mean irradiance, the algal-bacterial relationship was strengthened in the UVR-clear lake, where excreted organic carbon (EOC) rates exceeded the bacterial carbon demand (BCD). This did not occur in the UVR-opaque lake. The greater UVR damage to algae and bacteria and the weakening of their commensalistic interaction found in the UVR-opaque lake indicates that these ecosystems would be especially vulnerable to stressors related to global change. Thus, our findings may have important implications for the carbon cycle in oligotrophic lakes of the Mediterranean region.

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Evaluating how environmental stressor interactions influence ecosystem structure and functioning is critical to understanding the response of ecosystems to global change. We exposed a species-poor planktonic community to P pulses in the absence and presence of solar ultraviolet radiation (UVR). We used a field-mesocosm study in an oligotrophic mid-altitude lake to test the hypothesis that interaction between these factors affects the community's diversity and composition, but not its biomass-size spectrum, making this community resilient in terms of its C-transfer function. Our findings show that P pulses and UVR affected the relative abundance of different planktonic populations. The abundance of phytoplankton was enhanced strongly by P pulses and secondarily by UVR. The UVR Â P interaction affected only the smallest autotrophic species. Chlorococcal abundance increased, whereas chroococcal cyanobacteria decreased. However, UVR had a much more pronounced effect than P on the composition of microcrustaceans, and both factors interactively affected their biomass. Hence, the biomass-size spectrum was not resilient to the UVR Â P pulse interaction. The steepness of the slope increased under P-pulse conditions because zooplankton were not able to grow concomitantly with phytoplankton, as confirmed by the low zooplankton to phytoplankton biomass ratio (Z∶P). The observed planktonic changes under new foreseeable conditions that include an increase in UVR fluxes and P inputs might decouple the C cycle in inland oligotrophic lakes in the Mediterranean region.