Biological interactions underpin the functioning of marine ecosystems, be it via competition, predation, mutualism, or symbiosis processes. Microbial phototroph-heterotroph interactions propel the engine that results in the biogeochemical cycling of individual elements and are critical for understanding and modelling global ocean processes. Unfortunately, studies thus far have focused on exponentially-growing cultures in nutrient-rich media, meaning knowledge of such interactions under in situ conditions is rudimentary at best. Here, we performed long-term phototroph-heterotroph co-culture experiments under nutrient-amended and natural seawater conditions which showed that it is not the concentration of nutrients but rather their circulation that maintains a stable interaction and a dynamic system. Using the Synechococcus-Roseobacter interaction as a model phototroph-heterotroph case study we show that whilst Synechococcus is highly specialised for carrying out photosynthesis and carbon-fixation it relies on the heterotroph to re-mineralise the inevitably leaked organic matter making nutrients circulate in a mutualistic system. In this sense we challenge the general belief that marine phototrophs and heterotrophs compete for the same scarce nutrients and niche space, but instead suggest these organisms more likely benefit from each other because of their different levels of specialization and complementarity within long-term stable-state systems.

Mixotrophs combine photosynthesis with phagotrophy to cover their demands in energy and essential nutrients. This gives them a competitive advantage under oligotropihc conditions, where nutrients and bacteria concentrations are low. As the advantage for the mixotroph depends on light, the competition between mixo- and heterotrophic bacterivores should be regulated by light. To test this hypothesis, we incubated natural plankton from the ultra-oligotrophic Eastern Mediterranean in a set of mesocosms maintained at 4 light levels spanning a 10-fold light gradient. Picoplankton (heterotrophic bacteria (HB), pico-sized cyanobacteria, and small-sized flagellates) showed the fastest and most marked response to light, with pronounced predator-prey cycles, in the high-light treatments. Albeit cell specific activity of heterotrophic bacteria was constant across the light gradient, bacterial abundances exhibited an inverse relationship with light. This pattern was explained by light-induced top-down control of HB by bacterivorous phototrophic eukaryotes (PE), which was evidenced by a significant inverse relationship between HB net growth rate and PE abundances. Our results show that light mediates the impact of mixotrophic bacterivores. As mixo- and heterotrophs differ in the way they remineralize nutrients, these results have far-reaching implications for how nutrient cycling is affected by light.

We studied the effects of future climate change scenarios on plankton communities of a Norwegian fjord using a mesocosm approach. After the spring bloom, natural plankton were enclosed and treated in duplicates with inorganic nutrients elevated to pre-bloom conditions (N, P, Si; eutrophication), lowering of 0.4 pH units (acidification), and rising 3°C temperature (warming). All nutrient-amended treatments resulted in phytoplankton blooms dominated by chain-forming diatoms, and reached 13–16 μg chlorophyll (chl) a l−1. In the control mesocosms, chl a remained below 1 μg l−1. Acidification and warming had contrasting effects on the phenology and bloom-dynamics of autotrophic and heterotrophic microplankton. Bacillariophyceae, prymnesiophyceae, cryptophyta, and Protoperidinium spp. peaked earlier at higher temperature and lower pH. Chlorophyta showed lower peak abundances with acidification, but higher peak abundances with increased temperature. The peak magnitude of autotrophic dinophyceae and ciliates was, on the other hand, lowered with combined warming and acidification. Over time, the plankton communities shifted from autotrophic phytoplankton blooms to a more heterotrophic system in all mesocosms, especially in the control unaltered mesocosms. The development of mass balance and proportion of heterotrophic/autotrophic biomass predict a shift towards a more autotrophic community and less-efficient food web transfer when temperature, nutrients and acidification are combined in a future climate-change scenario. We suggest that this result may be related to a lower food quality for microzooplankton under acidification and warming scenarios and to an increase of catabolic processes compared to anabolic ones at higher temperatures.

Plankton biomass and composition in the pelagic zone of oceans is exposed to changes in availability of light and nutrients due to large-scale ocean circulation and water column stratification. We hypothesized that displacement of plankton from surface to deeper darker waters would not only favor heterotrophy over time, as previously suggested, but also first rapidly affect the level of mixotrophy and, consequently, overall microbial grazing in plankton food webs. To test this in an oligotrophic marine system we incubated Eastern Mediterranean water (from 10 m depth north of Crete in September 2010) in 2.8 m 3 mesocosms simulating two different light intensities at the sampling station, surface waters (ca. 10 m; mesocosms L1) and deeper layers (ca. 50-60 m; mesocosms L4). The biomass and abundance of the main planktonic groups were monitored either daily or every second day, depending on the group. Microzooplankton grazing rates and the contribution of mixotrophic feeding were estimated by a combination of dilution experiments and incubations with live fluorescently labeled algae (LFLA). Although no nutrients were added to the mesocosms the chlorophyll a increased during the first 2 days of the experiment in both treatments. This increase resulted from phytoplankton growth in the light L1-mesocosm (autotrophic biomass was ca. doubled in L1 compared to L4), but was mostly due to photoadaptation of the algae in the L4-mesocosm, as indicated by lower carbon to chlorophyll a ratios. By the end of the experiment, the total biomass of protozoan and metazoan grazers in L1 was ca. twofold higher than in L4. The microzooplankton responded within the first 24 h, showing different grazing activity in L1 than in L4. Microzooplankton grazing rates on total Chl a were similar in both treatments; however, phytoplankton instantaneous growth rates were higher in the more illuminated mesocosm. This resulted in a closer coupling between both rates