A mechanistic model for the uptake and elimination of ionizable organic chemicals at fish gills is presented. This model is a modification of a previous model for nonionizable organic chemicals that addressed the transport of chemical to and from gill surfaces in water and blood, diffusion of chemical across epithelial cells, and binding of chemical to components in water and blood. For ionizable chemicals, three additional processes are included. First, excretory products alter the pH at gill surfaces, affecting the relative amounts of neutral and ionized molecules compared with that in the bulk exposure water. Second, ionized molecules support chemical flux to and from epithelial cell membranes and help maintain high diffusion gradients of neutral molecules across these membranes, thereby contributing to uptake and elimination even if the membranes are impermeable to ionized molecules. Third, membrane barriers are not completely impermeable to ionized molecules, and even limited permeability can have appreciable effects on chemical flux. Approaches for model parameterization are discussed. Model-predicted relationships of uptake and elimination rates to exposure water pH, alkalinity, and chemical properties are presented and discussed in terms of model processes. The model is shown to predict important features of reported effects of pH on uptake rates of weak organic acids.
Effects of exposure-water pH on chemical uptake at rainbow trout (Oncorhynchus mykiss) gills were investigated for nine weakly acidic, chlorinated phenols with different ionization constants and hydrophobicities and for a moderately hydrophobic, nonionizable reference chemical (1,2,4-trichlorobenzene). Uptake rates for all chemicals varied little from pH 6.3 to 8.4, despite ionization of the chlorinated phenols ranging from less than 1 to greater than 99.9% among these pH values and chemicals. At pH 9.2, uptake rates were reduced substantially for the chlorinated phenols but not for the reference chemical. These results indicate greater bioavailability of neutral chemical forms but also considerable bioavailability of ionized forms that varies with pH. Three mechanisms were evaluated regarding such ionized chemical bioavailability. First, reduced pH at the gill surface causes net conversion of ionized molecules to more readily absorbed neutral molecules. This mechanism was tested by increasing exposure-water alkalinity, which increased gill surface pH and reduced uptake of the chlorinated phenols but not of the reference chemical. Magnitudes of these effects were close to predictions from a mathematical model for chemical exchange at fish gills that incorporated this mechanism. Second, ionized molecules contribute to uptake by maintaining high gradients of neutral molecules across epithelial membrane barriers, even if the barriers are impermeable to these ions. This mechanism was demonstrated to explain the similarity of uptake among pH values and chemicals at pH less than 8.4 and the degree to which uptake declined at pH 9.2. Third, membrane barriers can have some permeability to the ionized forms, but this was not important for the chemicals and conditions of the present study. Increased exposure-water pH also was demonstrated to increase elimination rates of these chemicals, which also was in accord with model expectations.
The relative importance of regional, watershed, and in-stream environmental factors on fish assemblage structure and function was investigated in western Lake Superior tributaries. We selected 48 second- and third-order watersheds from two hydrogeomorphic regions to examine fish assemblage response to differences in forest fragmentation, watershed storage, and a number of other watershed, riparian, and in-stream habitat conditions. Although a variety of regional, fragmentation, and storage-related factors had significant influences on the fish assemblages, water temperature appeared to be the single most important environmental factor. We found lower water temperatures and troutsculpin assemblages at lower fragmentation sites and higher temperatures and minnowsuckerdarter assemblages as storage increased. Factors related to riparian shading and flow separated brook trout streams from brown trout (Salmo trutta) rainbow trout (Oncorhynchus mykiss) streams. Functionally, fish assemblages at lower fragmentation sites were dominated by cold-water fishes that had low silt tolerance and preferred moderate current speeds, while fishes with higher silt tolerances, warmer temperature preferences, and weaker sustained swimming capabilities were most common at higher storage sites. Our results suggest that site-specific environmental conditions are highly dependent on regional- and watershed-scale characters and that a combination of these factors operates in concert to influence the structure and function of stream fish assemblages.
To facilitate extrapolation among watersheds, ecological risk assessments should be based on a model of underlying factors influencing watershed response, particularly vulnerability. We propose a conceptual model of landscape vulnerability to serve as a basis for watershed classification systems to predict resistance and resilience of aquatic ecosystems to hydrology-related stressors. Watershed area, storage capacity, channel slope, and soil permeability determine sensitivity of lotic systems to stressors associated with land-use activities that impact hydrologic regimes. Natural hydrologic disturbance regimes also influence the resilience of aquatic systems by selecting for life history strategies associated with rapid recolonization following disturbance. Variability in some of these physiographic driving factors can be partitioned by landscape classification schemes such as the U.S. Forest Service Ecological Unit Classification System, while others (watershed storage) may explain remaining variability within landscape units. We are conducting a comparative watershed study to examine simple and interactive effects of physiographic units, watershed storage (lakes ϩ wetlands), and land-clearing activities in watersheds surrounding the western arm of Lake Superior. Initial results for second-order watersheds indicate significant watershed class effects on baseflow water quality, percent motile biraphid diatom species in periphyton communities, habitat quality, and fish community integrity. Future studies have been designed to examine cumulative effects downstream.
To test a conceptual model of non-linear response of hydrologic regimes to watershed characteristics, we selected 48 secondand third-order study sites on the North and South Shores of western Lake Superior, MN (USA) using a random-stratified design based on hydrogeomorphic region, fraction mature forest, and fraction watershed storage (lakeCwetland area/watershed area). We calculated several commonly used hydrologic indices from discharge and velocity estimates, including daily flow indices, overall flood indices, low flow variables, and ratios or ranges of flow percentiles reflecting the nature of cumulative frequency distributions. Four principal components (PCs) explained 85.9 and 88.6% of the variation of flow metrics among second-and third-order stream sites, respectively. Axes of variation corresponded to a runoff vs. baseflow axis, flow variability, mean flow, and contrasts between flood duration and frequency. Analysis of velocity metrics for third-order streams yielded four PCs corresponding to mean or maximum velocity, Froude number, and inferred shear velocity, as well as spate frequencies vs. intervals associated with different velocity ranges.Using discriminant function analysis, we could discriminate among watershed classes based on region, mature forest, or watershed storage as a function of flow metrics. For second-order streams, median flow (Qs 50 ) increased as watershed storage increased. North Shore streams showed a more skewed distribution and greater spread of discharge values than did South Shore streams for both stream orders, while third-order North Shore streams exhibited a higher frequency of spates. Independent of regional differences, loss of mature forest increased the range of variation between baseflow and peak flows, and depressed baseflow. Consistent with our initial model for watershed classification, Classification and Regression Tree (CART) analysis confirmed significant thresholds of change in flow metrics averaging between 0.506 and 0.636 for fraction mature forest and between 0.180 and 0.258 for fraction watershed storage. q
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