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Mixotrophy, the combination of autotrophic and heterotrophic nutrition, is a common trophic strategy among unicellular eukaryotes in the ocean. There are a number of hypotheses about the conditions that select for mixotrophy, and field studies have documented the prevalence of mixotrophy in a range of environments. However, there is currently little evidence for how mixotrophy varies across environmental gradients, and whether empirical patterns support theoretical predictions. Here I synthesize experiments that have quantified the abundance of phototrophic, mixotrophic, and heterotrophic nanoflagellates, to ask whether there are broad patterns in the prevalence of mixotrophy (relative to pure autotrophy and heterotrophy), and to ask whether observed patterns are consistent with a trait-based model of trophic strategies. The data suggest that mixotrophs increase in abundance at lower latitudes, while autotrophs and heterotrophs do not, and that this may be driven by increased light availability. Both mixotrophs and autotrophs increase greatly in productive coastal environments, while heterotrophs increase only slightly. These patterns are consistent with a model of resource competition in which nutrients and carbon can both limit growth and mixotrophs experience a trade-off in allocating biomass to phagotrophy vs. autotrophic functions. Importantly, mixotrophy is selected for under a range of conditions even when mixotrophs experience a penalty for using a generalist trophic strategy, due to the synergy between photosynthetically derived carbon and prey-derived nutrients. For this reason mixotrophy is favored relative to specialist strategies by increased irradiance, while at the same time increased nutrient supply increases the competitive ability of mixotrophs against heterotrophs.

Temperature strongly affects phytoplankton growth rates, but its effect on communities and ecosystem processes is debated. Because phytoplankton are often limited by light, temperature should change community structure if it affects the traits that determine competition for light. Furthermore, the aggregate response of phytoplankton communities to temperature will depend on how changes in community structure scale up to bulk rates. Here, we synthesize experiments on 57 phytoplankton species to analyze how the growth-irradiance relationship changes with temperature. We find that light-limited growth, light-saturated growth, and the optimal irradiance for growth are all highly sensitive to temperature. Within a species, these traits are co-adapted to similar temperature optima, but light-limitation reduces a species’ temperature optimum by ~5°C, which may be an adaptation to how light and temperature covary with depth or reflect underlying physiological correlations. Importantly, the maximum achievable growth rate increases with temperature under light saturation, but not under strong light limitation. This implies that light limitation diminishes the temperature sensitivity of bulk phytoplankton growth, even though community structure will be temperature-sensitive. Using a database of primary production incubations, we show that this prediction is consistent with estimates of bulk phytoplankton growth across gradients of temperature and irradiance in the ocean. These results indicate that interactions between temperature and resource limitation will be fundamental for explaining how phytoplankton communities and biogeochemical processes vary across temperature gradients and respond to global change.

Nutrient utilization traits can be used to link the ecophysiology of phytoplankton to population dynamic models and the structure of communities across environmental gradients. Here we analyze a comprehensive literature compilation of four traits: maximum nutrient uptake rate; the half-saturation constant for nutrient uptake; the minimum subsistence quota, measured for nitrate and phosphate; and maximum growth rate. We also use these traits to analyze two composite traits, uptake affinity and scaled uptake affinity. All traits tend to increase with cell volume, except for scaled uptake affinity and maximum growth rate, which tend to decline with cell volume. Most scaling relationships are the same for freshwater and marine species, although important differences exist. Most traits differ on average between major taxa, but between-taxon variation is nearly always due to between-taxon variation in volume. There is some evidence for between-trait correlations that could constrain trait evolution, but these correlations are difficult to disentangle from correlation driven by cell volume. These results should enhance the parameterization of models that use size or taxonomic group to structure physiological variation in phytoplankton communities.

Light-dependent growth of phytoplankton is a fundamental process in marine ecosystems, but we lack a comprehensive view of how light utilization traits vary across genotypes and species, and how this variation is structured by cell size, taxonomy, and environmental gradients. Here, we compile 308 growth-irradiance experiments performed on 119 species of marine phytoplankton from all major functional groups, and characterize growth-irradiance relationships in terms of the initial slope of the growth-irradiance curve (a), the optimal irradiance above which growth declines (I opt), and the maximum growth rate (μ max). We find that a declines with increasing cell size, although cell size appears to be a weak constraint on this trait. There are significant differences across taxa in α and μ max, with dinoflagellates, raphidophytes, and diazotrophs having the lowest values for both traits, and Phaeocystis spp. and diatoms having relatively high values. I opt does not vary among taxonomic groups, and all traits exhibit large variation within most groups. Open-ocean isolates tend to have higher α, lower I opt, and lower μ max than coastal isolates, implying adaptation to low light and low productivity. The three traits are correlated across species such that α and I opt are negatively related while μ max is positively correlated with both of these traits. There is some evidence that high a carries a cost of high N demand even when nitrogen (not light) is limiting. The results elucidate contrasting light-related ecological strategies across phytoplankton and should help improve the parameterization of major functional groups in biogeochemical models.

Phytoplankton are key players in the global carbon cycle, contributing about half of global primary productivity. Within the phytoplankton, functional groups (characterized by distinct traits) have impacts on other major biogeochemical cycles, such as nitrogen, phosphorus and silica. Changes in phytoplankton community structure, resulting from the unique environmental sensitivities of these groups, may significantly alter elemental cycling from local to global scales. We review key traits that distinguish major phytoplankton functional groups, how they affect biogeochemistry and how the links between community structure and biogeochemical cycles are modelled. Finally, we explore how global environmental change will affect phytoplankton communities, from the traits of individual species to the relative abundance of functional groups, and how that, in turn, may alter biogeochemical cycles. Synthesis. We can increase our mechanistic understanding of the links between the community structure of primary producers and biogeochemistry by focusing on traits determining functional group responses to the environment (response traits) and their biogeochemical functions (effect traits). Identifying trade‐offs including allometric and phylogenetic constraints among traits will help parameterize predictive biogeochemical models, enhancing our ability to anticipate the consequences of global change.

Small eukaryotic phytoplankton are major contributors to global primary production and marine biogeochemical cycles. Many taxa are thought to be mixotrophic, but quantitative studies of phagotrophy exist for very few. In addition, little is known about consumers of Prochlorococcus , the abundant cyanobacterium at the base of oligotrophic ocean food webs. Here we describe thirty-nine new phytoplankton isolates from the North Pacific Subtropical Gyre (Station ALOHA), all flagellates ~2–5 µm diameter, and we quantify their ability to graze Prochlorococcus . The mixotrophs are from diverse classes (dictyochophytes, haptophytes, chrysophytes, bolidophytes, a dinoflagellate, and a chlorarachniophyte), many from previously uncultured clades. Grazing ability varied substantially, with specific clearance rate (volume cleared per body volume) varying over ten-fold across isolates and six-fold across genera. Slower grazers tended to create more biovolume per prey biovolume consumed. Using qPCR we found that the haptophyte Chrysochromulina was most abundant among the isolated mixotrophs at Station ALOHA, with 76–250 cells mL −1 across depths in the upper euphotic zone (5–100 m). Our results show that within a single ecosystem the phototrophs that ingest bacteria come from many branches of the eukaryotic tree, and are functionally diverse, indicating a broad range of strategies along the spectrum from phototrophy to phagotrophy.

Trade-offs among functional traits are essential for explaining community structure and species coexistence. While two-way trade-offs have been investigated in many systems, higher-dimensional trade-offs remain largely hypothetical. Here we demonstrate a three-way trade-off between cell size and competitive abilities for nitrogen and phosphorus in marine and freshwater phytoplankton. At a given cell size, competitive abilities for N and P are negatively correlated, but as cell size increases, competitive ability decreases for both nutrients. The relative importance of the two trade-off axes appears to be environment dependent, suggesting different selective pressures: freshwater phytoplankton separate more along the N vs. P competition axis, and marine phytoplankton separate more along the nutrient competition vs. cell size axis. Our results demonstrate the multidimensional nature of key trade-offs among traits and suggest that such trade-offs may drive species interactions and structure ecological communities.

Resource pulses are infrequent, large-magnitude, and short-duration events of increased resource availability. They include a diverse set of extreme events in a wide range of ecosystems, but identifying general patterns among the diversity of pulsed resource phenomena in nature remains an important challenge. Here we present a meta-analysis of resource pulse–consumer interactions that addresses four key questions: (1) Which characteristics of pulsed resources best predict their effects on consumers? (2) Which characteristics of consumers best predict their responses to resource pulses? (3) How do the effects of resource pulses differ in different ecosystems? (4) What are the indirect effects of resource pulses in communities? To investigate these questions, we built a data set of diverse pulsed resource–consumer interactions from around the world, developed metrics to compare the effects of resource pulses across disparate systems, and conducted multilevel regression analyses to examine the manner in which variation in the characteristics of resource pulse–consumer interactions affects important aspects of consumer responses. Resource pulse magnitude, resource trophic level, resource pulse duration, ecosystem type and subtype, consumer response mechanisms, and consumer body mass were found to be key explanatory factors predicting the magnitude, duration, and timing of consumer responses. Larger consumers showed more persistent responses to resource pulses, and reproductive responses were more persistent than aggregative responses. Aquatic systems showed shorter temporal lags between peaks of resource availability and consumer response compared to terrestrial systems, and temporal lags were also shorter for smaller consumers compared to larger consumers. The magnitude of consumer responses relative to their resource pulses was generally smaller for the direct consumers of primary resource pulses, compared to consumers at greater trophic distances from the initial resource pulse. In specific systems, this data set showed both attenuating and amplifying indirect effects. We consider the mechanistic processes behind these patterns and their implications for the ecology of resource pulses.

Mixotrophic eukaryotes are important bacterivores in oligotrophic open oceans, but their significance as grazers in more nutrient‐rich waters is less clear. Here, we investigated the bacterivory partition between mixotrophs and heterotrophs in a productive, estuary‐influenced coastal region in the East China Sea. We found ubiquitous, actively feeding phytoplankton populations and taxa with mixotrophic potential by identifying ingestion of fluorescent prey surrogate and analyzing community 18S rRNA gene amplicons. Potential and active mixotrophs accounted for 10–63% of the total eukaryotic community and 17–69% of bacterivores observed, respectively, contributing 6–48% of estimated in situ bacterivory. The much higher mixotroph fitness outside of the turbid plume was potentially driven by increased light and decreased nutrient availability. Our results suggest that, although heterotrophs dominated overall in situ bacterivory, mixotrophs were abundant and important bacterivores in this low‐latitude mesotrophic coastal region.

Viruses infecting aquatic microbes vary immensely in size, but the ecological consequences of virus size are poorly understood. Here we used a unique suite of diverse phytoplankton strains and their viruses, all isolated from waters around Hawai'i, to assess whether virus size affects the suppression of host populations. We found that small viruses of diverse genome type (3–24 kb genome size, 23–70 nm capsid diameter) have very similar effects on host populations, suppressing hosts less strongly and for a shorter period of time compared to large double‐stranded DNA viruses (214–1380 kb, 112–386 nm). Suppressive effects of larger viruses were more heterogeneous, but most isolates reduced host populations by many orders of magnitude, without recovery over the ~ 25‐d experiments. Our results suggest that disparate lineages of viruses may have ecological consequences that are predictable in part based on size, and that ecosystem impacts of viral infection may vary with the size structure of the viral community.