Divvying up the pie: Tissue nutrient content is related to its parasite load.

The nutrient content of host resources can influence the abundance of parasites within an ecosystem, but linking specific nutrients in a host to the abundance of different parasite taxa remains a challenge. Here, we work to forge this link by quantifying the relationship between the nutrient content of specific infection sites and the abundance of multiple parasite taxa within the digestive tract of largemouth bass (Micropterus salmoides) collected from the Mississippi River. To generate a mechanistic understanding of these relationships, we tested four basic predictions: (1) the nutrient content of different host tissues (infection sites) varies within and across hosts, (2) the nutrient content of parasite genera differs from that of their host tissue(s), (3) the nutrient content of parasite genera differ from one another and (4) the nutrient content of host tissues is related to the nutrient content and abundance of parasite genera. We found support for each of these predictions. We found stoichiometric differences between the digestive tissues we examined. We also found that across hosts, intestine and pyloric caeca C:N ratios increased and %N decreased with fish condition factor. Both of the actively feeding parasitic genera we measured had lower C:N ratios compared to both their host tissue and other encysted/non-reproductive genera, suggesting the potential for N limitation of these parasites in the intestines or pyloric caeca of hosts. Consistent with this possibility, we found that the total number of actively feeding parasitic worms in the pyloric caeca increased with that tissue's N:P ratio (but was not related to host condition factor). Our results suggest that parasites encounter significant variation in nutrient content within and across hosts and that this variation may influence the abundance of actively feeding parasites. This work highlights the need for additional empirical comparisons of parasite stoichiometry across tissues and individual hosts.

Partitioning macroscale and microscale ecological processes using covariate-driven non-stationary spatial models.

Ecological inference requires integrating information across scales. This integration creates a complex spatial dependence structure that is most accurately represented by fully non-stationary models. However, ecologists rarely use these models because they are difficult to estimate and interpret. Here, we facilitate the use of fully non-stationary models in ecology by improving the interpretability of a recently developed non-stationary model and applying it to improve our understanding of the spatial processes driving lake eutrophication. We reformulated a model that incorporates non-stationary correlation by adding environmental predictors to the covariance function, thereby building on the intuition of mean regression. We created ellipses to visualize how data at a given site correlate with their surroundings (i.e., the range and directionality of underlying spatial processes). We applied this model to describe the spatial dependence structure of variables related to lake eutrophication across two different regions: a Midwestern United States region with highly agricultural landscapes, and a Northeastern United States region with heterogeneous land use. For the Midwest, increases in forest cover increased the homogeneity of the residual spatial structure of total phosphorus, indicating that macroscale processes dominated this nutrient's spatial structure. Conversely, high forest cover and baseflow reduced the spatial homogeneity of chlorophyll a residuals, indicating that microscale processes dominated for chlorophyll a in the Midwest. In the Northeast, increases in urban land use and baseflow decreased the homogeneity of phosphorus concentrations indicating the dominance of microscale processes, but none of our covariates were strongly associated with the residual spatial structure of chlorophyll a. Our model showed that the spatial dependence structure of environmental response variables shifts across space. It also helped to explain this structure using ecologically relevant covariates from different scales whose effects can be interpreted intuitively. This provided novel insight into the processes that lead to eutrophication, a complex and pervasive environmental issue.

Nutrient availability affects the prevalence of a microsporidian parasite.

Defining the relationship between nutrients and parasitism is complicated by shifts in host physiology and population density, which can both mediate the effects of host diet on parasites and vice versa. We examined the relationship between nutrient availability and an abundant parasite capable of both horizontal and vertical transmission (Hamiltosporidium tvaerminnensis) of a planktonic crustacean, Daphnia magna, in rock pools on Baltic Sea Skerry islands. We found that the relative availability of nutrients directly affected infection prevalence; parasite prevalence was higher in pools with higher particulate N:P ratios. Infection prevalence was not related to Daphnia population densities. A complementary experiment that examined host responses to an N:P gradient in mesocosms indicated that high N:P ratios can increase spore load in the hosts. We surmise that high N:P food increases Daphnia feeding rate, which increases their contact with parasite spores and leads to higher prevalence and more intense infections. We found no direct evidence that parasite-induced changes in host nutrient use affected nutrient dynamics in pools. However, the relationship between diet N:P and the parasite's prevalence and load is consistent with previously documented patterns of this parasite's effect on host nutrient use. Taken together, this study suggests that high N:P ratios in food may benefit the parasite in multiple ways and could create environments that favour horizontal transmission over vertical transmission for parasites capable of both transmission routes. If so, nutrient limitation could have long-term consequences for host-parasite evolution.

Quantifying the effects of surface conveyance of treated wastewater effluent on groundwater, surface water, and nutrient dynamics in a large river floodplain.

Restoration and reconnection of floodplain systems provide multiple societal and ecosystem benefits, while providing municipalities the opportunity to attempt alternative approaches to maintain infrastructure protection and function. In some restored floodplains, treated wastewater effluent discharge is redirected over land instead of directly into rivers to allow natural flow and infiltration, to facilitate restoration designs such as levee setback, and to provide additional freshwater to floodplain ecosystems. However, indirect discharge of treated effluent over land may pose risks to surface and groundwater when pollutants like excess nutrients enter the floodplain and undergo transformation. We investigated the consequences for groundwater and surface water quality when effluent was redirected as open water channels over a floodplain surface. In this study, seasonal floodplain nutrient concentrations in groundwater and surface water were observed for more than 5 years as a floodplain and wastewater treatment plant underwent a major restoration project that included river-floodplain reconnection with levee setback and redirection of effluent discharge from a river channel to open flow across the restored floodplain. Nutrient loading to the surrounding floodplain groundwater and surface water was observed, but based on measures of hydrological connectivity, groundwater flow paths, and biogeochemistry, nutrients from the effluent moved within the floodplain with minimal effect to the surrounding floodplain water quality. We did not find evidence of substantial additional processing that could replace advanced nutrient treatment in this system, however we did observe evidence of diverse nutrient processes that may support enhanced retention if treatment channels were designed to enhance these processes. We suggest that indirect discharge of high quality treated effluent in a restored floodplain is a viable alternative to direct discharge into a river when groundwater flow directs that discharge to habitats where minimal nutrient sensitivity is expected.

Exploited and excreting: parasite type affects host nutrient recycling.

Parasite-induced changes in the nutrient balance of hosts could alter the availability of nutrients in ecosystems by changing consumer-driven nutrient recycling. While these effects on host nutrient use are mediated by host physiology, they likely depend on characteristics of the parasite and host diet quality. We examined this possibility by measuring nutrient release rates of uninfected Daphnia and conspecifics infected by two microparasites (the bacterium Pasteuria ramosa and the microsporidium Hamiltosporidium tvaerminnensis) from daphnid hosts fed food that varied in phosphorus content. We found that infection type and diet affected host nutrient release rates, but the strength of these effects varied among parasite treatments. To improve our understanding of these effects, we examined whether two separate aspects of host exploitation (parasite-induced reductions in host fecundity and parasite load) could account for variation in Daphnia nutrient release, ingestion, and elemental ratios caused by our infection and diet treatments. Regardless of whether we compared individuals across infection type or diet treatment, Daphnia fecundity described variation in multiple aspects of host nutrient use better than infection, diet, or spore load. Our results suggest that parasite-induced changes in host nutrient use are both parasite and diet specific, and that host fecundity could be a useful parameter for predicting the magnitude and direction of these changes.

Does infection tilt the scales? Disease effects on the mass balance of an invertebrate nutrient recycler.

While parasites are increasingly recognized as important components of ecosystems, we currently know little about how they alter ecosystem nutrient availability via host-mediated nutrient cycling. We examined whether infection alters the flow of nutrients through hosts and whether such effects depend upon host diet quality. To do so, we compared the mass specific nutrient (i.e., nitrogen and phosphorus) release rates, ingestion rates, and elemental composition of uninfected Daphnia to those infected with a bacterial parasite, P. ramosa. N and P release rates were increased by infection when Daphnia were fed P-poor diets, but we found no effect of infection on the nutrient release of individuals fed P-rich diets. Calculations based on the first law of thermodynamics indicated that infection should increase the nutrient release rates of Daphnia by decreasing nutrient accumulation rates in host tissues. Although we found reduced nutrient accumulation rates in infected Daphnia fed all diets, this reduction did not increase the nutrient release rates of Daphnia fed the P-rich diet because infected Daphnia fed this diet ingested nutrients more slowly than uninfected hosts. Our results thus indicate that parasites can significantly alter the nutrient use of animal consumers, which could affect the availability of nutrients in heavily parasitized environments.