Global warming has revitalized interest in the relationship between body size and temperature, proposed by Bergmann's rule 150 years ago, one of the oldest manifestations of a ‘biogeography of traits’. We review biogeographic evidence, results from clonal cultures and recent micro‐ and mesocosm experiments with naturally mixed phytoplankton communities regarding the response of phytoplankton body size to temperature, either as a single factor or in combination with other factors such as grazing, nutrient limitation, and ocean acidification. Where possible, we also focus on the comparison between intraspecific size shifts and size shifts resulting from changes in species composition. Taken together, biogeographic evidence, community‐level experiments and single‐species experiments indicate that phytoplankton average cell sizes tend to become smaller in warmer waters, although temperature is not necessarily the proximate environmental factor driving size shifts. Indirect effects via nutrient supply and grazing are important and often dominate. In a substantial proportion of field studies, resource availability is seen as the only factor of relevance. Interspecific size effects are greater than intraspecific effects. Direct temperature effects tend to be exacerbated by indirect ones, if warming leads to intensified nutrient limitation or copepod grazing while ocean acidification tends to counteract the temperature effect on cell size in non‐calcifying phytoplankton. We discuss the implications of the temperature‐related size trends in a global‐warming context, based on known functional traits associated with phytoplankton size. These are a higher affinity for nutrients of smaller cells, highest maximal growth rates of moderately small phytoplankton (ca. 10² µm³), size‐related sensitivities for different types of grazers, and impacts on sinking rates. For a phytoplankton community increasingly dominated by smaller algae we predict that: (i) a higher proportion of primary production will be respired within the microbial food web; (ii) a smaller share of primary production will be channeled to the classic phytoplankton – crustacean zooplankton – fish food chain, thus leading to decreased ecological efficiency from a fish‐production point of view; (iii) a smaller share of primary production will be exported through sedimentation, thus leading to decreased efficiency of the biological carbon pump.

While the isolated responses of marine phytoplankton to climate warming and to ocean acidification have been studied intensively, studies on the combined effect of both aspects of Global Change are still scarce. Therefore, we performed a mesocosm experiment with a factorial combination of temperature (9 and 15°C) and pCO₂ (means: 439 ppm and 1040 ppm) with a natural autumn plankton community from the western Baltic Sea. Temporal trajectories of total biomass and of the biomass of the most important higher taxa followed similar patterns in all treatments. When averaging over the entire time course, phytoplankton biomass decreased with warming and increased with CO₂ under warm conditions. The contribution of the two dominant higher phytoplankton taxa (diatoms and cryptophytes) and of the 4 most important species (3 diatoms, 1 cryptophyte) did not respond to the experimental treatments. Taxonomic composition of phytoplankton showed only responses at the level of subdominant and rare species. Phytoplankton cell sizes increased with CO₂ addition and decreased with warming. Both effects were stronger for larger species. Warming effects were stronger than CO₂ effects and tended to counteract each other. Phytoplankton communities without calcifying species and exposed to short-term variation of CO₂ seem to be rather resistant to ocean acidification.

Shrinking of body size has been proposed as one of the universal responses of organisms to global climate warming. Using phytoplankton as an experimental model system has supported the negative effect of warming on body-size, but it remains controversial whether the size reduction under increasing temperatures is a direct temperature effect or an indirect effect mediated over changes in size selective grazing or enhanced nutrient limitation which should favor smaller cell-sizes. Here we present an experiment with a factorial combination of temperature and nutrient stress which shows that most of the temperature effects on phytoplankton cell size are mediated via nutrient stress. This was found both for community mean cell size and for the cell sizes of most species analyzed. At the highest level of nutrient stress, community mean cell size decreased by 46% per °C, while it decreased only by 4.7% at the lowest level of nutrient stress. Individual species showed qualitatively the same trend, but shrinkage per °C was smaller. Overall, our results support the hypothesis that temperature effects on cell size are to a great extent mediated by nutrient limitation. This effect is expected to be exacerbated under field conditions, where higher temperatures of the surface waters reduce the vertical nutrient transport.

Decreasing body size has been suggested as the third universal biological response to global warming after latitudinal/altitudinal range shifts and shifts in phenology. Size shifts in a community can be the composite result of intraspecific size shifts and of shifts between differently sized species. Metabolic explanations for the size shifts dominate in the literature but top down effects, i.e. intensified size-selective consumption at higher temperatures, have been proposed as alternative explanation. Therefore, we performed phytoplankton experiments with a factorial combination of warming and consumer type (protist feeding mainly on small algae vs. copepods mainly feeding on large algae). Natural phytoplankton was exposed to 3 (1^st experiment) or 4 (2^nd experiment) temperature levels and 3 (1^st experiment: nano-, microzooplankton, copepods) or 2 (2^nd experiment: microzooplankton, copepods) types of consumers. Size shifts of individual phytoplankton species and community mean size were analyzed. Both, mean cell size of most of the individual species and mean community cell size decreased with temperature under all grazing regimes. Grazing by copepods caused an additional reduction in cell size. Our results reject the hypothesis, that intensified size selective consumption at higher temperature would be the dominant explanation of decreasing body size. In this case, the size reduction would have taken place only in the copepod treatments but not in the treatments with protist grazing (nano- and microzooplankton).

Cyanobacterial blooms are the most important and best studied type of harmful algal blooms in fresh waters and brackish coastal seas. We here review how and to which extent they resist grazing by zooplankton, how zooplankton responds to cyanobacterial blooms and how these effects are further transmitted to fish. Size, toxicity and poor nutritional value are widespread mechanisms of grazing defense by cyanobacteria. In some cases, defenses are inducible, in some they are obligate. However, to some extent zooplankton overcome grazing resistance, partly after evolutionary adaptation. Cyanotoxins are also harmful to fish and may cause fish kills. However, some fish species feed on Cyanobacteria, are able to reduce their abundance, and grow on a cyanobacterial diet. While reduced edibility for crustacean zooplankton tends to elongate the food chain from primary producers to fish, direct feeding by fish tends to shorten it. The few available comparative studies relating fish yield to nutrients or phytoplankton provide no indication that cyanobacteria should reduce the ratio fish production: primary production.

Phytoplankton cell or colony sizes range from <1 µm to several cm, i.e. 4–5 orders of magnitude in linear dimensions, which is roughly equivalent to the log-size span within terrestrial vegetation. It is commonplace to assume that smaller phytoplankton have an advantage in growth related traits while larger ones are more resistant to losses. However, the current state of literature calls for a more differentiated view. It is still controversial, whether smaller phytoplankton have higher maximal growth rates (µ_max) or if there is a peak of µ_max at intermediate size (10² µm³ cell volume). Smaller phytoplankton have an advantage in nutrient acquisition at low concentrations while larger phytoplankton have an advantage in utilizing nutrient pulses and exploiting vertical gradients. At equal density, larger phytoplankton experience bigger sinking losses. Small phytoplankton (10⁵ µm³) in lake phytoplankton and the absence of large (>10³ µm³) green algae in marine plankton. Overall, size is one of the most important traits for the performance of phytoplankton, but it is overly simplistic to equate small size with metabolic advantages.