Individual vs. population plastic responses to elevated CO₂, nutrient availability, and heterogeneity: A microcosm experiment with co-occurring species

We conducted an experiment to evaluate the plastic phenotypic responses of individuals, growing under intra-specific competition, and populations of three co-occurring grassland species (Lolium perenne, Plantago lanceolata, and Holcus lanatus) to joint variations in atmospheric CO₂ partial pressure (P _CO2; 37.5 vs. 70 Pa), nutrient availability (NA; 40 vs. 120 mg N added as organic material), and the spatial pattern of nutrient supply (SH; homogeneous vs. heterogeneous nutrient supply). At both the population and individual levels, the aboveground biomass of the three species significantly increased when the nutrients were heterogeneously supplied. Significant two- (SH × NA) and three-term (P _CO2 × NA × SH) interactions determined the response of traits measured on populations (aboveground biomass and below: aboveground biomass ratio, BAR) and individuals (aboveground biomass and specific leaf area). The combination of a high SH and NA elicited the highest plasticity of aboveground biomass in populations and individuals of the three species evaluated, and of BAR in Holcus. Soil heterogeneity and elevated P _CO2 elicited the highest plasticity in the SLA of Plantago and Lolium individuals. Our results show that populations, and not only individuals, respond to soil heterogeneity in a plastic way, and that plastic responses to elevated P _CO2 are complex since they vary across traits and species, and are influenced by the availability of nutrients and by their spatial distribution. They also emphasize the importance of soil heterogeneity as a modulator of plant responses to global change drivers.