Although species commonly modify habitats and thereby influence ecosystem structure and function, the factors governing the ecological importance of these modifications are not well understood. Pacific salmon have repeatedly been shown to positively influence the abundance of benthic biota by annually transferring large quantities of nutrients from marine systems to the nutrient-poor freshwaters in which they spawn. Conversely, other studies have demonstrated that salmon can negatively influence the abundance of freshwater biota, an effect attributed to bioturbation during upstream migration and nest construction. The factors determining which of these contrasting ecological effects predominates are unknown, including how human activities, such as land use, influence ecological responses to salmon. We sampled a key basal food resource, sediment biofilm, in seven southeast Alaskan streams impacted to varying degrees by timber harvest. Biofilm abundance (measured as chlorophyll a and ash-free dry mass) was positively related to timber-harvest intensity prior to salmon arrival. However, during the salmon run, an inverse relationship emerged of more abundant biofilm in less-harvested watersheds. Among-stream variability in biofilm response to salmon was largely explained by sediment particle size, which was larger in less-harvested watersheds. Collectively, these results suggest that, by altering stream sediment size, timber harvest transformed the dominant effect of salmon from nutrient enrichment to physical disturbance, thus modifying nutrient linkages between marine and freshwater ecosystems.

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Using nutrient-diffusing substrata (NDS) in seven streams in southeast Alaska, USA, we tested whether (i) nutrient limitation of autotrophic and heterotrophic biofilms was alleviated by salmon resource subsidies, and (ii) whether the degree of alleviation could be predicted by environmental variables. Before salmon spawners arrived, autotrophic biofilms were nitrogen (N)-limited, or co-limited by N and phosphorus (P), whereas heterotrophic biofilms were either P-limited, or co-limited by N and P. Combined N and P amendments resulted in a 2.6-fold increase in biofilm chlorophyll a, and a 3.2-fold increase in community respiration. After salmon arrived, autotroph nutrient limitation was alleviated in six of the seven streams. Heterotrophs still exhibited nutrient limitation in six streams, but most streams shifted from co-limitation to P-limitation. Nutrient-diffusing substrata amended with salmon tissue indicated that salmon could also be an important source of organic carbon for biofilms. Autotrophs responded less to N and P amendments as streamwater ammonium concentration increased with the arrival of salmon. For heterotrophs, ammonium concentration and N:P ratio best predicted changes in response following the arrival of salmon. We provide the first direct evidence that biofilm nutrient limitation can be alleviated by salmon spawners in nutrient-poor streams.

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SCoC is patient-centered, outcomes-driven, value-based approach for hospital-wide surgical patient safety. The principles of this value paradigm are adaptable to other hospitals as demonstrated in our longitudinal study in 3 hospital systems, and the initial experience of CoC suggests that this model will have benefit beyond surgical hospital cohort.

1. Pacific salmon (Oncorhynchus spp.) deliver marine-derived nutrients to the streams in which they spawn and die, and these resource subsidies can increase the abundance of stream biota. In strong contrast, physical disturbance from salmon spawning activity can reduce the abundance of benthic organisms. Previous experimental designs have not established the relative effects of these two contrasting processes on stream organisms during a salmon run. 2. We combined manipulative and observational field studies to assess the degree of nutrient enrichment, physical disturbance, and the net effect of salmon on the abundance of benthic periphyton. Related salmon-mediated processes were also evaluated for benthic macroinvertebrates. Mesh exclosures (2 · 2 m plots) prevented salmon from disturbing areas of the stream channel, which were compared with areas to which salmon had access. Sampling was conducted both before and during the late-summer spawning run of pink (O. gorbushca) and chum (O. keta) salmon. 3. Streamwater nitrogen and phosphorus concentrations increased sharply with the onset of the salmon run, and highly significant positive relationships were observed between the numbers of salmon present in the stream and these dissolved nutrients. Before the salmon run, periphyton biomass (as chlorophyll a) and total macroinvertebrate abundance were very similar between control and exclosure plots. During the salmon run, exclosures departed substantially from controls, suggesting significant disturbance imparted on benthic biota. 4. Comparing exclosures before and during the salmon run enabled us to estimate the effects of salmon in the absence of direct salmon disturbance. This 'nutrient enrichment potential' was significant for periphyton biomass, as was a related index for macroinvertebrate abundance (although enhanced invertebrate drift into exclosures during the salmon run could also have been important). Interestingly, however, the net effect of salmon, evaluated by comparing control plots before and during the salmon run, was relatively modest for both periphyton and macroinvertebrates, suggesting that nutrient enrichment effects were largely offset by disturbance. 5. Our results illustrate the importance of isolating the specific mechanisms via which organisms affect ecosystems, and indicate that the relative magnitude of salmon nutrient enrichment and benthic disturbance determines the net effect that these ecologically important fish have on stream ecosystems.

1. Variation in resource subsidies can create or reinforce heterogeneity in recipient ecosystems. Related activities of organisms delivering resource subsidies, such as ecosystem engineering by Pacific salmon spawners (Oncorhynchus spp.), also alter heterogeneity. We studied whether heterogeneity in stream environmental conditions and spawner abundances were reflected in the net ecological effects of salmon (i.e. enrichment by resource subsidies and disturbance by ecosystem engineering) on benthic biofilm. 2. We sampled seven Southeast Alaska streams over 3 years, both before and during the salmon run. In each stream and year, stream environmental characteristics and their influence on responses of benthic biofilm [mean and coefficient of variation of chlorophyll a (chl a), ash-free dry mass (AFDM) and autotrophic index (AFDM:chl a)] to spawners were assessed. 3. Streams and periods before and during the salmon run were distinct based on their environmental characteristics. The responses of most biofilm metrics to spawners were streamand year-specific, suggesting that the ecological effect of spawners ranged from net enrichment to net disturbance depending on the stream or year studied. The environmental context, especially temperature, large wood, and sediment size, explained >50% of biofilm variability during the run, but <30% over the entire study, suggesting that salmon can alter environmental constraints. 4. Precision of biofilm estimates improved by increasing either the number of streams or the number of years sampled (i.e. spatial or temporal replication). However, combining data from different North Pacific Rim ecoregions inflated the confidence interval as compared with a single ecoregion, indicating the importance of regional environmental contexts for net salmon effects. 5. Our results suggest that biofilm responses to salmon can vary greatly, even within a single ecoregion, and that environmental conditions can modify net salmon effects. Consequently, generalisations about biofilm responses across the native range of salmon may be challenging.