Environmentally friendly microbeads to model the dispersal of particulates in aquatic systems.

The transport of particulate matter including the gametes, larvae and propagules of reproducing organisms and other organic matter involved in nutrient/contaminant transport are important processes, yet there are few environmentally friendly methods available to examine dispersal empirically. Herein we report on the development and application of a biodegradable and non-toxic physical model, based on alginate microbeads with modifiable size, density (ρ), and colour for use in dispersal studies. Specifically, the microbeads were designed to model the size and ρ of parasitic juvenile freshwater mussels (Unionidae; ρ = 1200 kg m-3), which undergo dispersal upon excystment from fish hosts. We released the juvenile-mussel and neutrally buoyant microbeads (ρ = 1000 kg m-3) in a local river and captured them in drift nets downstream. The concentration of microbeads declined with downstream distance, but neutrally buoyant microbeads were transported farther. Analysis of microbead capture rates could be described using the patterns of several mathematical models (negative exponential, power, and turbulent transport), which were consistent with the reported dispersal of mussel larvae and other benthic macroinvertebrates. These results support the use of alginate microbeads in dispersal studies, because their environmentally friendly and customizable properties offer improvements over non-biodegradable alternatives.

Threats to freshwater mussels: The interactions of water temperature, velocity and total suspended solids on ecophysiology and growth.

Freshwater unionid mussels are ecosystem engineers that are highly endangered in part because of land-use changes that have altered their habitat and negatively impacted their ecophysiology. The environmental factors that affect mussels do not act alone and may be better understood using a multiple-stressor approach. We examine how changes in water temperature, turbidity (total suspended solids; TSS) and water velocity affected the clearance rates (CR), oxygen consumption rates (OC), and resultant scope for growth (SFG) of Lampsilis siliquoidea in laboratory flow chamber experiments. The CR, OC and SFG of L. siliquoidea increased with acclimation temperature and velocity, and decreased with TSS concentration and acute temperature exposure, although these responses were more complicated when factors were combined. The primary factor affecting CR and OC varied with acclimation temperature, with warmer temperature and high TSS leading to strong declines in clearance rates. A worst-case scenario would involve a summer season where temperatures and TSS loads are above-average, and water velocities are either below- or above- average, which are likely under increased frequencies of storm, flood, or drought events due to climate change. Conservation measures should focus on protecting aquatic systems during these times and also use a multistressor approach to determine how environmental factors interact in efforts to protect and recover freshwater mussel populations.

Increases in Great Lake winds and extreme events facilitate interbasin coupling and reduce water quality in Lake Erie.

Climate change affects physical and biogeochemical processes in lakes. We show significant increases in surface-water temperature (~ 0.5 °C decade-1; > 0.2% year-1) and wave power (> 1% year-1; the transport of energy by waves) associated with atmospheric phenomena (Atlantic Multidecadal Oscillation and Multivariate El Niño/Southern Oscillation) in the month of August between 1980 and 2018 in the Laurentian Great Lakes. A pattern in wave power, in response to extreme winds, was identified as a proxy to predict interbasin coupling in Lake Erie. This involved the upwelling of cold and hypoxic (dissolved oxygen < 2 mg L-1) hypolimnetic water containing high total phosphorus concentration from the seasonally stratified central basin into the normally well-mixed western basin opposite to the eastward flow. Analysis of historical records indicate that hypoxic events due to interbasin exchange have increased in the western basin over the last four decades (43% in the last 10 years) thus affecting the water quality of the one of the world's largest freshwater sources and fisheries.

Collector Motion Affects Particle Capture in Physical Models and in Wind Pollination.

Particle capture is important for ecological processes in aquatic and terrestrial ecosystems. The current model is based on a stationary collector for which predictions about capture efficiency (η; flux of captured particles ∶ flux of particles) are based on the collector flow environment (i.e., collector Reynolds number, Rec; inertial force ∶ viscous force). This model does not account for the movement of collectors in nature. We examined the effect of collector motion (transverse and longitudinal to the flow) on η using a cylindrical model in the lab and the grass species Phleum pratense in the field. Collector motion increased η (up to 400% and 20% in the lab and field, respectively) and also affected the spatial distribution of particles on collectors, especially at low Rec. The effect was greatest for collectors moving transversely at large magnitude, which encountered more particles with higher relative momentum. These results, which differ from the stationary model, can be predicted by considering both Rec and the particle dynamics given by the Stokes number (Stk; particle stopping distance ∶ collector radius) and helped to resolve an existing controversy about pollination mechanisms. Collector motion should be considered in wind pollination and other ecological processes involving particle capture.

Suspended solid concentration reduces feeding in freshwater mussels.

We examined the effect of TSS concentration on the clearance rates (CR) of newly transformed juvenile and adult Lampsilis fasciola, L. siliquoidea, Ligumia nasuta, and Villosa iris, as increased total suspended solids (TSS) are thought to interfere with feeding processes. Mussel CR were measured in aerated (or swirled for juveniles) chambers at TSS concentrations up to 15mgL-1 for laboratory-transformed juveniles, and up to 100mgL-1 for adult mussels. The CR of one-week old animals increased with TSS concentration, likely due to ontological differences in feeding (pedal vs. suspension feeding) and gill development, but CR decreased monotonically with TSS concentration in older animals (two-, three- and four-week old juveniles). The CR of adult mussels were significantly lower at TSS concentrations ≥8mgL-1, which represented a threshold in CR. Although this threshold occurred at similar concentrations across the four species, the decline in CR was largest in L. fasciola (46% compared to no-TSS control), and smallest in V. iris (21%). Differences among species are likely related to differences in the TSS and substrate found in their source rivers given that greater decline occurred for species in rivers with relatively lower TSS. The decrease in CR as TSS increased is consistent across marine and freshwater bivalves, at both juvenile and adult stages. The decrease in feeding was five times greater in juvenile compared to adult bivalves, which indicates how the vulnerability to environmental stressors differ across life stages. These results demonstrate that TSS reduces suspension feeding rates in freshwater unionids, therefore TSS should be managed to ensure their survival.

Loss of reproductive output caused by an invasive species.

We investigated whether Neogobius melanostomus, an invader of biodiversity 'hot-spots' in the Laurentian Great Lakes region, facilitates or inhibits unionid mussel recruitment by serving as a host or sink for their parasitic larvae (glochidia). Infestation and metamorphosis rates of four mussel species with at-risk (conservation) status (Epioblasma torulosa rangiana, Epioblasma triquetra, Lampsilis fasciola and Villosa iris) and one common species (Actinonaias ligamentina) on N. melanostomus were compared with rates on known primary and marginal hosts in the laboratory. All species successfully infested N. melanostomus, but only E. triquetra, V. iris and A. ligamentina successfully metamorphosed into juveniles, albeit at very low rates well below those seen on even the marginal hosts. Neogobius melanostomus collected from areas of unionid occurrence in the Grand and Sydenham rivers (Ontario, Canada) exhibited glochidial infection rates of 39.4% and 5.1%, respectively, with up to 30 glochidia representing as many as six unionid species per fish. A mathematical model suggests that N. melanostomus serve more as a sink for glochidia than as a host for unionids, thereby limiting recruitment success. This represents a novel method by which an invasive species affects a native species.

The effect of bottom roughness on scalar transport in aquatic ecosystems: implications for reproduction and recruitment in the benthos.

Bottom roughness can influence gamete and larval transport in benthic organisms. For example the ratio of the roughness spacing (λ) and roughness height (k) determines the type of roughness flow regime created in two dimensional (2D) flows: λ/k<8 results in skimming flow; λ/k~8 results in wake interference flow; and λ/k>8 results in isolated roughness flow. Computational fluid dynamic modeling (COMSOL K-ε) was used to examine the effect of roughness geometry (e.g., a gradient in angularity provided by square, triangular and round 2D bottom roughness elements) on the prediction of roughness flow regime using biologically relevant λ/k ratios. In addition, a continuously released scalar (a proxy for gametes and larvae) in a coupled convection-diffusion model was used to determine the relationship among roughness geometry, λ/k ratios, and scalar transport (relative scalar transport, RT=ratio of scalar measured downstream in a series of roughness elements placed in tandem). The modeled roughness flow regimes fit closely with theoretical predictions using the square and triangle geometries, but the round geometry required a lower λ/k ratio than expected for skimming flow. Relative transport of the scalar was consistent with the modeled flow regimes, however significant differences in RT were found among the roughness flows for each geometry, and significantly lower RT values were observed for skimming flow in the round geometry. The λ/k ratio provides an accurate means of classifying flow in and around the roughness elements, whereas RT indicates the nature of scalar transport and retention. These results indicate that the spatial configuration of bottom roughness is an important determinant of gamete/larval transport in terms of whether the scalar will be retained among roughness elements or transported downstream.

Turbulence-induced resonance vibrations cause pollen release in wind-pollinated Plantago lanceolata L. (Plantaginaceae).

In wind pollination, the release of pollen from anthers into airflows determines the quantity and timing of pollen available for pollination. Despite the ecological and evolutionary importance of pollen release, wind-stamen interactions are poorly understood, as are the specific forces that deliver pollen grains into airflows. We present empirical evidence that atmospheric turbulence acts directly on stamens in the cosmopolitan, wind-pollinated weed, Plantago lanceolata, causing resonant vibrations that release episodic bursts of pollen grains. In laboratory experiments, we show that stamens have mechanical properties corresponding to theoretically predicted ranges for turbulence-driven resonant vibrations. The mechanical excitation of stamens at their characteristic resonance frequency caused them to resonate, shedding pollen vigorously. The characteristic natural frequency of the stamens increased over time with each shedding episode due to the reduction in anther mass, which increased the mechanical energy required to trigger subsequent episodes. Field observations of a natural population under turbulent wind conditions were consistent with these laboratory results and demonstrated that pollen is released from resonating stamens excited by small eddies whose turnover periods are similar to the characteristic resonance frequency measured in the laboratory. Turbulence-driven vibration of stamens at resonance may be a primary mechanism for pollen shedding in wind-pollinated angiosperms. The capacity to release pollen in wind can be viewed as a primary factor distinguishing animal- from wind-pollinated plants, and selection on traits such as the damping ratio and flexural rigidity may be of consequence in evolutionary transitions between pollination systems.

The effect of natural dissolved organic carbon on the acute toxicity of copper to larval freshwater mussels (glochidia).

The present study examined the effect of dissolved organic carbon (DOC), both added and inherent, on Cu toxicity in glochidia, the larvae of freshwater mussels. Using incremental additions of natural DOC concentrate and reconstituted water, a series of acute copper toxicity tests were conducted. An increase in DOC from 0.7 to 4.4 mg C/L resulted in a fourfold increase (36-150 µg Cu/L) in the 24-h median effective concentration (EC50) and a significant linear relationship (r² = 0.98, p = 0.0008) between the DOC concentration and the Cu EC50 of Lampsilis siliquoidea glochidia. The ameliorating effect of added DOC on Cu toxicity was confirmed using a second mussel species, the endangered (in Canada) Lampsilis fasciola. The effect of inherent (i.e., not added) DOC on Cu toxicity was also assessed in eight natural waters (DOC 5-15 mg C/L). These experiments revealed a significant relationship between the EC50 and the concentration of inherent DOC (r² = 0.79, p = 0.0031) with EC50s ranging from 27 to 111 µg Cu/L. These laboratory tests have demonstrated that DOC provides glochidia with significant protection from acute Cu toxicity. The potential risk that Cu poses to mussel populations was assessed by comparing Cu and DOC concentrations from significant mussel habitats in Ontario to the EC50s. Although overall mean Cu concentration in the mussel's habitat was well below the acutely toxic level given the concentration of DOC, episodic Cu releases in low DOC waters may be a concern for the recovery of endangered freshwater mussels. The results are examined in the context of current Cu water quality regulations including the U.S. Environmental Protection Agency's (U.S. EPA) biotic ligand model.

Sensitivity of the glochidia (larvae) of freshwater mussels to copper: assessing the effect of water hardness and dissolved organic carbon on the sensitivity of endangered species.

The assessment of the potential impact of waterborne contaminants on imperilled freshwater mussels is needed. Acute copper toxicity was assessed in a standardized soft water (hardness 40-48 mg CaCO(3)equivalents L(-1)) using the larvae (glochidia) from three common and six (Canadian) endangered mussel species. The resulting 24h EC50s ranged from 7 to 36 microg Cu L(-1), with the EC50s of two endangered species <10 microg Cu L(-1). Acute copper sensitivity was also determined in Ptychobranchus fasciolaris, a species that employs conglutinates (packets of glochidia) in its reproductive strategy. Conglutinates were found to provide significant protection from acute copper exposure as the EC50 of the encased glochidia was more than four-fold higher than freed glochidia (72.6 microg Cu L(-1) vs. 16.3 microg Cu L(-1)). The glochidia from two endangered species, Epioblasma triquetra and Lampsilis fasciola, were used to examine the influence of water hardness and dissolved organic carbon (DOC) on copper sensitivity. Exposures in moderately-hard water (165 mg CaCO(3) L(-1)) demonstrated that an increase in water hardness resulted in a significant reduction in copper sensitivity. For example, in L. fasciola the 24 h EC50s were 17.6 (14.2-22.6) microg Cu L(-1) and 50.4 (43.5-60.0) microg Cu L(-1) in soft water and moderately-hard water, respectively. The addition of DOC (as Aldrich Humic Acid) also resulted in a significant decrease in Cu sensitivity, such that a 10-fold increase in the EC50 of E. triquetra was observed when the reconstituted soft water was augmented with 1.6 mg DOC L(-1). To determine if current water quality regulations for copper would protect all glochidia, the USEPA's Biotic Ligand Model (BLM) was used to derive water quality criteria for these exposures. While BLM-derived criteria for the soft water exposures indicate that protection would be marginal for the sensitive endangered species, the criteria derived for the DOC exposures suggest that the natural complexity of most natural waters in Southern Ontario (Canada) will provide glochidia with protection from acute copper exposure.

The interaction of CO2 concentration and spatial location on O2 flux and mass transport in the freshwater macrophytes Vallisneria spiralis and V. americana.

The biology of aquatic organisms determines the maximum rates of physiological processes, but the mass transport of nutrients determines the nominal rates at which these processes occur. Maximum O(2) flux (P(max)) at 17.1 mmol m(-3) CO(2) was higher for the leaves of the freshwater macrophyte Vallisneria spiralis [P(max)=0.013+/-0.001 mmol m(-2) s(-1) (g(chla+b) m(-2))(-1) (mean +/- s.e.m.)] than for the closely related species, Vallisneria americana [P(max)=0.008+/-0.001 mmol m(-2) s(-1) (g(chla+b) m(-2))(-1)]. The O(2) flux saturated at freestream velocities >4.5+/-1.2 cm s(-1) and was spatially invariant for both species. However, a tenfold decrease in CO concentration to 1.71 mmol m(-3) changed the nature of the relationship between O(2) flux and spatial location along the leaf surface, and reduced the O(2) flux of V. spiralis to values similar to V. americana. The O(2) flux [P(max)=0.007+/-0.001 mmol m(-2) s(-1) (g(chla+b) m(-2))(-1)] saturated at the upstream location (i.e. 1 cm from the leading edge of the leaf) but was found to increase linearly with freestream velocity [slope=0.057+/-0.011 mmol m(-2) s(-1) (g(chla+b) m(-2))(-1) (m s(-1))(-1)] at the downstream location (i.e. 7 cm from the leading edge) at freestream velocities >1.8+/-0.9 cm s(-1). Conversely, mass transfer rates did not vary with CO(2) concentration, and were characteristic of a laminar concentration boundary layer at the upstream location and a turbulent concentration boundary layer at the downstream location. Rates of mass transfer measured directly from O(2) profiles were not predicted by theoretical values based on hydrodynamic measurements. Moreover, the concentration boundary layer thickness (delta(CBL)) values measured directly from O(2) profiles were 48+/-2% and 21+/-1% of the predicted theoretical delta(CBL) values at the upstream and downstream locations, respectively. It is evident that physiological processes involving mass transport are coupled and vary in space. Mass transport investigations of biological systems based solely on hydrodynamic measurements need to be interpreted with caution.