Experimental test of phytoplankton competition for nutrients and light in poorly mixed water columns

A recent theory of the vertical distribution of phytoplankton considers how interacting niche construction processes such as resource depletion, behavior, and population dynamics contribute to spatial heterogeneity in the aquatic environment. In poorly mixed water columns with opposing resource gradients of nutrients and light, theory predicts that a species should aggregate at a single depth. This depth of aggregation, or biomass maximum, should change through time due to depletion of available resources. In addition, the depth of the aggregation should be deeper under low amounts of nutrient loading and shallower under higher amounts of nutrient loading. Theory predicts total biomass to exhibit a saturating relationship with nutrient supply. A surface biomass maximum limited by light and a deep biomass maximum limited by nutrients or co-limited by nutrients and light is also predicted by theory. To test this theory, we used a motile phytoplankton species (Chlamydomonas reinhardtii ) growing in cylindrical plankton towers. In our experiment, the resource environment was strongly modified by the movement, self-shading, nutrient uptake, and growth of the phytoplankton. Supporting predictions, we routinely observed a single biomass maximum at the surface throughout the course of the experiment and at equilibrium under higher nutrient loading. However, at equilibrium, low nutrient loading led to a non-distinct biomass maximum with the population distributed over most of the water column instead of the distinct subsurface peak predicted by theory. Also supporting predictions, total biomass across water columns was positively related to nutrient supply but saturating at high nutrient supply conditions. Further supporting predictions, we also found evidence of light limitation for a surface biomass maximum and nutrient limitation for the deep biomass when no surface maximum was present. In addition, the light level leaving the bottom of the water column declined through time as the phytoplankton grew and was negatively related to nutrient loading. Nutrients were strongly depleted where biomass was present by the end of the experiment. This experimental study shows that the vertical distribution of phytoplankton may be driven by intraspecific resource competition in space.