Ecosystem complexity explains the scale-dependence of ammonia toxicity on macroinvertebrates

The toxic effect of unionized ammonia (NH₃) on aquatic organisms is receiving increasing attention due to the excessive nitrogen discharge to various surface waters. Researches have suggested the scale-dependence of NH₃ toxicity, being lower in field than under lab conditions. Such scale-dependence of toxicity is a big challenge to water quality criteria setting as the results solely from lab tests might not apply to natural ecosystems. Therefore, it is necessary to explore the underlying mechanism to understand the difference of toxicity across various spatial scales. In this study, we used the widely distributed gastropod Bellamya aeruginosa as the test animal and performed two 192-h microcosm experiments. Each experiment included a control and an ammonia addition treatment: N0(LC₅₀) & N+(LC₅₀), N0(LC₁₀₀) & N+(LC₁₀₀) (96-h LC₅₀ = 0.8 mg NH₃N/L, 96-h LC₁₀₀ = 18.1 mg NH₃N/L). Besides water-only, three potential key components (food, sediment, and submersed macrophytes) were included in the various treatments to mimic different complexity levels of aquatic ecosystems (Water, Water + Food, Water + Sediment, Water + Sediment + Macrophytes). The results showed that: 1) food directly improved the survival and growth of gastropods under expected lethal concentration of ammonia (96-h concentration of NH₃N = LC₂₀ of the 96-h acute test); 2) sediment and macrophyte quickly decreased the ammonia concentration, mainly by sediment adsorption and macrophyte uptake, to alleviate the ammonia stress to gastropods and permitted them to survive and grow under expected lethal concentration of ammonia (96-h concentration of NH₃N = LC₁₀∼LC₂₀ of the 96-h acute test); 3) sediment and macrophyte also provided additional food for gastropods; 4) under extremely high ammonia stress (i.e., 96-h LC₁₀₀, food was left uneaten and macrophyte died, and gastropods could, therefore, not be released from ammonia stress. Our results demonstrate that under moderate ammonia stress, the introduction of extra ecosystem elements (food, sediment, and macrophytes) significantly improved the survival and growth of gastropods, mainly by enhancing their tolerance and by quickly decreasing the NH₃ concentration and thus toxicity. However, these introduced elements had little effect at very high concentration of ammonia (i.e., 96-h LC₁₀₀). Our findings add to the understanding of the reasons behind the previous observed scale-dependent toxicity of NH₃ on aquatic organisms and contribute to better decisions on the role of NH₃ in relation to water quality management.