  Resource competition in heterogeneous environments is still an unresolved
problem of theoretical ecology. In this article I analyze competition between
two phytoplankton species in a deep water column, where the distributions of
main resources (light and a limiting nutrient) have opposing gradients and
co-limitation by both resources causes a deep biomass maximum. Assuming that
the species have a trade-off in resource requirements and the water column is
weakly mixed, I apply the invasion threshold analysis (Ryabov and Blasius 2011)
to determine relations between environmental conditions and phytoplankton
composition. Although species deplete resources in the interior of the water
column, the resource levels at the bottom and surface remain high. As a result,
the slope of resources gradients becomes a new crucial factor which, rather
than the local resource values, determines the outcome of competition. The
value of resource gradients nonlinearly depend on the density of consumers.
This leads to complex relationships between environmental parameters and
species composition. In particular, it is shown that an increase of both the
incident light intensity or bottom nutrient concentrations favors the best
light competitors, while an increase of the turbulent mixing or background
turbidity favors the best nutrient competitors. These results might be
important for prediction of species composition in deep ocean.
