Light, nutrients, and food-chain length constrain planktonic energy transfer efficiency across multiple trophic levels.

The efficiency of energy transfer through food chains [food chain efficiency (FCE)] is an important ecosystem function. It has been hypothesized that FCE across multiple trophic levels is constrained by the efficiency at which herbivores use plant energy, which depends on plant nutritional quality. Furthermore, the number of trophic levels may also constrain FCE, because herbivores are less efficient in using plant production when they are constrained by carnivores. These hypotheses have not been tested experimentally in food chains with 3 or more trophic levels. In a field experiment manipulating light, nutrients, and food-chain length, we show that FCE is constrained by algal food quality and food-chain length. FCE across 3 trophic levels (phytoplankton to carnivorous fish) was highest under low light and high nutrients, where algal quality was best as indicated by taxonomic composition and nutrient stoichiometry. In 3-level systems, FCE was constrained by the efficiency at which both herbivores and carnivores converted food into production; a strong nutrient effect on carnivore efficiency suggests a carryover effect of algal quality across 3 trophic levels. Energy transfer efficiency from algae to herbivores was also higher in 2-level systems (without carnivores) than in 3-level systems. Our results support the hypothesis that FCE is strongly constrained by light, nutrients, and food-chain length and suggest that carryover effects across multiple trophic levels are important. Because many environmental perturbations affect light, nutrients, and food-chain length, and many ecological services are mediated by FCE, it will be important to apply these findings to various ecosystem types.

Nutrient cycling by fish supports relatively more primary production as lake productivity increases.

Animals can be important in nutrient cycling in particular ecosystems, but few studies have examined how this importance varies along environmental gradients. In this study we quantified the nutrient cycling role of an abundant detritivorous fish species, the gizzard shad (Dorosoma cepedianum), in reservoir ecosystems along a gradient of ecosystem productivity. Gizzard shad feed mostly on sediment detritus and excrete sediment-derived nutrients into the water column, thereby mediating a cross-habitat translocation of nutrients to phytoplankton. We quantified nitrogen and phosphorus cycling (excretion) rates of gizzard shad, as well as nutrient demand by phytoplankton, in seven lakes over a four-year period (16 lake-years). The lakes span a gradient of watershed land use (the relative amounts of land used for agriculture vs. forest) and productivity. As the watersheds of these lakes became increasingly dominated by agricultural land, primary production rates, lake trophic state indicators (total phosphorus and chlorophyll concentrations), and nutrient flux through gizzard shad populations all increased. Nutrient cycling by gizzard shad supported a substantial proportion of primary production in these ecosystems, and this proportion increased as watershed agriculture (and ecosystem productivity) increased. In the four productive lakes with agricultural watersheds (>78% agricultural land), gizzard shad supported on average 51% of phytoplankton primary production (range 27-67%). In contrast, in the three relatively unproductive lakes in forested or mixed-land-use watersheds (>47% forest, <52% agricultural land), gizzard shad supported 18% of primary production (range 14-23%). Thus, along a gradient of forested to agricultural landscapes, both watershed nutrient inputs and nutrient translocation by gizzard shad increase, but our data indicate that the importance of nutrient translocation by gizzard shad increases more rapidly. Our results therefore support the hypothesis that watersheds and gizzard shad jointly regulate primary production in reservoir ecosystems.

Interactive effects of light and nutrients on phytoplankton stoichiometry.

The stoichiometric composition of autotrophs can vary greatly in response to variation in light and nutrient availability, and can mediate ecological processes such as C sequestration, growth of herbivores, and nutrient cycling. We investigated light and nutrient effects on phytoplankton stoichiometry, employing five experiments on intact phytoplankton assemblages from three lakes varying in productivity and species composition. Each experiment employed two nutrient and eight irradiance levels in a fully factorial design. Light and nutrients interactively affected phytoplankton stoichiometry. Thus, phytoplankton C:N, C:P, and N:P ratios increased with irradiance, and slopes of the stoichiometric ratio versus irradiance relationships were steeper with ambient nutrients than with nutrients added. Our results support the light-nutrient hypothesis, which predicts that phytoplankton C:nutrient ratios are functions of the ratio of available light and nutrients; however, we observed considerable variation among lakes in the expression of this relationship. Phytoplankton species diversity was positively correlated with the slopes of the C:N and C:P versus irradiance relationships, suggesting that diverse assemblages may exhibit greater flexibility in the response of phytoplankton nutrient stoichiometry to light and nutrients. The interactive nature of light and nutrient effects may render it difficult to generate predictive models of stoichiometric responses to these two factors. Our results point to the need for future studies that examine stoichiometric responses across a wide range of phytoplankton communities.