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Elevated allochthonous inputs of organic matter are increasingly recognized as a driver of ecosystem change in lakes, particularly when concurrent with eutrophication. Evaluation of lakes in a nutrient-color paradigm (i.e., based on total phosphorus and true color) enables a more robust approach to research and management. To assess temporal and spatial patterns in nutrient-color status for U.S. lakes and associated food web attributes, we analyzed the U.S. Environmental Protection Agency's National Lakes Assessment (NLA) data. With 1000+ lakes sampled in 2007 and 2012 in a stratified random sampling design, the NLA enables rigorous assessment of lake condition across the continental U.S. We demonstrate that many U.S. lakes are simultaneously experiencing eutrophication and brownification to produce an abundance of "murky" lakes. Overall, "blue" lakes decreased by ~ 18% (46% of lakes in 2007 to 28% in 2012) while "murky" lakes increased by almost 12% (24% of lakes in 2007 to 35.4% in 2012). No statistical differences were observed in the proportions of "green" or "brown" lakes. Regionally, murky lakes significantly increased in the Northern Appalachian, Southern Plains, and Xeric ecoregions. Murky lakes exhibited the highest epilimnetic chlorophyll a concentrations, cyanobacterial densities, and microcystin concentrations. Total zooplankton biomass was also highest in murky lakes, primarily due to increased rotifer and copepod biomass. However, zooplankton : phytoplankton biomass ratios were low, suggesting reduced energy transfer to higher trophic levels. These results emphasize that many lakes in the U.S. are simultaneously "greening" and "browning", with potentially negative consequences for water quality and food web structure.

We examined seasonal and spatial patterns in dissolved organic carbon (DOC) and chromophoric dissolved organic matter (CDOM) in the Chowan River watershed, North Carolina, a blackwater river which discharges into the second largest estuary in the United States, the Albemarle–Pamlico Estuarine System. From April 2008 to May 2010, DOC concentration did not significantly vary across seasons (range 7.69–30.39 mg L−1); however, CDOM molecular size and aromaticity increased throughout the spring, decreased during the summer and fall, and remained relatively low in the winter. Spectral slope ratios suggested microbial processing of CDOM in the spring and photodegradation of CDOM in the summer and fall. Spatially, DOC and CDOM concentrations were similar in the mainstem and at the mouths of two tributaries, Bennetts Creek and Wiccacon River, but were significantly higher upstream on the tributaries. DOC concentration was positively correlated with CDOM absorbance coefficients at 254 and 350 nm; however, these optical proxies explained only ~60 % of the variance. DOC and CDOM absorption loads to the Albemarle Sound ranged from 2.63 × 1010 g year−1 and 9.84 × 1010 m2 year−1, respectively, in a dry year and 7.9 × 1010 g year−1 and 2.2 × 1011 m2 year−1, respectively, in a wet year, which are comparable to non-blackwater rivers with larger watersheds. Blackwater rivers may therefore represent “hotspots” in coastal carbon chemistry, with seasonal variations in the quality and quantity of DOC and CDOM influencing estuarine food web dynamics and net ecosystem metabolism.

Organic matter fluxes and food web interactions in lakes depend on the abilities of heterotrophic microbial communities to access and degrade organic matter, a process that begins with extracellular hydrolysis of high molecular weight substrates. In order to determine whether patterns of enzymatic hydrolysis vary among shallow lakes of different trophic status, we investigated the hydrolysis of six specific organic macromolecules (polysaccharides) in the spring and late summer in four adjacent shallow lakes of eutrophic, oligotrophic, and dystrophic status in coastal North Carolina, USA. The spectrum of enzyme activities detected differed strongly between lakes, with all six polysaccharides hydrolyzed in West Mattamuskeet in May, while only two substrates were hydrolyzed in Lake Phelps in August/September. Differences in the spectrum of enzyme activities, and therefore the capabilities of heterotrophic microbial communities, were likely driven by variations among lakes in primary productivity patterns, sediment–water interactions, and/or water chemistry. Our data represent a first step towards a better understanding of carbon substrate availability and differences in carbon cycling pathways in shallow lakes of different trophic status.

The intensity and spectral composition of visible light are known to influence fish predation on zooplankton. However, in clear-water systems, ultraviolet (UV) radiation (UVR) may also influence fish predation, directly through UV-enhanced foraging, indirectly through alterations in predator-prey overlap, or in a combination of the two. Here we test the hypothesis that UVR facilitates fish predation on zooplankton in an oligotrophic lake. Experiments were conducted in 2.2-m-long UV-transparent or UV-blocking columns suspended in the epilimnion. Zooplankton consumption by young-of-the-year (YOY) largemouth bass (Micropterus salmoides) was compared in the presence and absence of UVR, and the vertical distributions of prey were quantified across light and fish treatments. In addition, a separate experiment was conducted to examine zooplankton vertical responses to UV exposure vs. fish kairomones. Overall, YOY predation on zooplankton was higher in the presence than in the absence of UVR, particularly on diaptomid copepods. This result was only partially explained by zooplankton migratory behaviors, suggesting UV-enhanced searching capabilities in YOY bass. Furthermore, diaptomids displayed a stronger vertical behavioral response to fish kairomones than to UVR whereas Daphnia exhibited a stronger response to UVR.

In transparent, thermally stratified lakes, ultraviolet radiation (UV) and temperature are among the primary factors influencing diel vertical migration (DVM) of zooplankton. It is not well known how behavioral responses will vary across a wide range of depths with steep gradients in UV, photosynthetically active radiation (PAR), and temperature. Our objective was to determine the relative importance of UV and temperature on DVM of Daphnia catawba and Leptodiaptomus minutus in a high UV, thermally stratified lake. Using UV-transparent (+UV) and UV-opaque (–UV) columns suspended within the mixed layer and metalimnion, we found that both species were generally shallower in the –UV than in the +UV columns. Daphnia catawba responded negatively to UV, even below the 1% depth for 320 nm, whereas L. minutus responded to UV only within the mixed layer and upper metalimnion. Daphnia catawba did not migrate in the –UV in the mixed layer, but migrated upwards in the –UV in the deep part of the metalimnion, indicating a temperature-induced rather than phototactic response. At night, both species migrated upwards in the metalimnion and were evenly distributed in the mixed layer. These results indicate that in transparent lakes, UV may constrain some zooplankton to cooler, suboptimal temperatures, which may compromise fitness.

Previous studies indicate that leachates derived from plants grown under elevated CO 2 are more recalcitrant and consequently reduce bacterial productivity. Here bacteria were cultured on organic leachates derived from trembling aspen (Populus tremuloides) and white willow (Salix alba) grown under ambient (360 ppm) and elevated (720 ppm) CO 2 concentrations and exposed and unexposed to simulated solar radiation. In treatments not exposed to light, determined numbers of colony forming units (CFU mL -1 ) were higher on ambient CO 2 than elevated CO 2 leachates in both plant species. After exposure to simulated sunlight, the number of CFU mL -1 decreased on the ambient CO 2 leachates and increased on the elevated leachates, particularly the aspen. These results demonstrate that environmental parameters, mediated by climate change, may have both positive and negative effects on the growth of bacteria.

Ultraviolet (UV) photoreceptors have been reported in a wide variety of freshwater and marine organisms, suggesting that UV vision is prominent in aquatic ecosystems. However, its adaptive significance remains speculative. The present study tested whether the foraging of juvenile bluegill (Lepomis macrochirus) is enhanced in the presence of UV radiation (UVR). Laboratory feeding trials were conducted in a laminar flow tank in which L. macrochirus juveniles between 2.3 and 3.5 cm standard length were fed the cladoceran Daphnia magna. Sighting and striking distances, as well as capture success, were measured in the presence and absence of UVR. Mean sighting and striking distances and capture success did not differ significantly between the two light treatments. There were also no significant differences in the frequency distributions of sighting and striking distance. These results suggest that UV vision may not be used to enhance foraging in L. macrochirus on Daphnia within the size class tested.

Zooplankton taxa exhibit varying tolerances to ultraviolet radiation (UVR), with rotifers and copepods tending to be more UV tolerant than cladocerans, regardless of lake UV transparency. On the basis of these differences, we hypothesized that UV avoidance behavior would be greatest in zooplankton with low UV tolerance, particularly in high-UV systems. Both downward and upward movements of zooplankton were examined in cylindrical acrylic columns that either transmitted full sunlight or blocked UV-B and short wavelength UV-A radiation. Columns were suspended vertically in the surface waters of a high- and low-UV lake at varying times of day and cloud cover. In the high-UV lake, the cladoceran Daphnia catawba displayed a distinct avoidance of the surface waters in the UV+ treatment but often showed a preference for the surface in the UV– treatment. Copepods were less responsive to UVR with calanoids displaying a somewhat greater UV avoidance than cyclopoids. In the low-UV lake, no behavioral differences were observed among UV treatments. These results suggest that UVR may influence the vertical distribution and habitat partitioning of certain zooplankton taxa in high-UV lakes while other factors, such as predation and food limitation, may be more important in low-UV lakes.

Recently, small-scale field experiments have demonstrated that exposure to UV radiation can invoke downward migrations in UV-sensitive zooplankton; however, questions remain as to zooplankton responses at the whole-lake scale. Here a three-year database on three lakes of varying UV transparency was used to examine the relationship between UV exposure and the seasonal vertical distribution and abundance of several zooplankton species. We hypothesized that if UV radiation is an important ecological force influencing the vertical distribution and seasonal abundance of zooplankton, the strength of a given species' response to UV would be inversely proportional to its UV tolerance. UV habitat did have a significant effect on zooplankton distribution in taxa with low UV tolerance compared to taxa with high UV tolerance. Cladocerans, which are the least UV-tolerant of three major groups, tended to avoid the surface waters of the highest UV lake during periods of high UV. More UV-tolerant groups, rotifers and copepods, showed little or no evidence for avoidance of high UV habitats. Among lakes, the relationship between UV avoidance and taxon was evident only in the highest UV lake. These data are consistent with the hypothesis that UV influences zooplankton community dynamics in low DOC, UV-transparent lakes.

In nature most organisms have to manage conflicting demands of food gathering, predator avoidance, and finding a favorable abiotic environment (oxygen, temperature, etc.) in order to maximize their fitness. In the vertical water column of lakes with high solar ultraviolet radiation (UV) and invertebrate predators, zooplankton face two particularly strong and conflicting selective pressures. During daylight hours invertebrate predators often induce an upward vertical migration of zooplankton prey while potentially damaging UV forces a downward migration. We used 2.2 m long columns suspended vertically in a lake to conduct 2x2 factorial experiments to examine patterns of depth selection behavior by zooplankton in the presence and absence of both the invertebrate predator Chaoborus and UV. We hypothesized that Chaoborus and UV both affect the distribution of zooplankton and a combination of both factors would lead to a narrowing of depth distribution. We found that when Chaoborus were present zooplankton tended to be distributed at shallower depths in the columns, while in the presence of UV they exhibited a deeper distribution. Chaoborus themselves were always found near the bottom of the columns regardless of the UV treatment. Simultaneous exposure to predators and UV resulted in a peak of zooplankton (especially Daphnia catawba) distribution at intermediate depths. In a significant number of cases, depth range was narrowed in response to Chaoborus, UV, or both.

Damaging solar radiation was one of the first factors proposed to explain zooplankton diel vertical migrations (DVM), yet other factors such as temperature, food, and especially predation have typically been more widely studied and identified as the primary factors inducing DVM. This is true in spite of the fact that recent experiments have shown that ambient levels of solar ultraviolet radiation (UVR) are potentially lethal to zooplankton and negative phototactic behavior has been demonstrated in the laboratory. Here we present the first evidence from in situ experiments demonstrating that UVR wavelengths influence the vertical swimming behavior of zooplankton under full spectrum solar radiation. Cylindrical acrylic columns that transmitted or blocked UVR were used to examine the behavioral responses of Daphnia pulicaria to UVR. Experiments were conducted in the epilimnion of a high-UV system at midday and in late afternoon. In both experiments, a large proportion of individuals migrated downward in the presence of UVR, whereas those in the UVR-shielded treatments tended to remain closer to the surface.

Solar ultraviolet radiation (UVR) has been demonstrated to have damaging effects on zooplankton, but little is known about what factors influence UVR tolerance in nature. Here we examined the relationship between UVR tolerance (the sum of photopro- tection and photorepair processes) and zooplankton taxon, body size, and source lake UVR transparency. Zooplankton of various sizes and taxa from lakes of different UV transparency were exposed to different intensities of a constant artificial UVR source. UVR tolerance was expressed as the UVR dose at which 50% mortality was observed for a given species. Smaller zooplankton species showed a uniformly high UVR tolerance, while larger zoo- plankton varied in their UVR tolerance both among and within species. The smaller rotifers, Keratella in particular, showed a high UVR tolerance while the larger, more transparent rotifer (Asplanchna) showed an intermediate UVR tolerance. Both cyclopoid and calanoid copepod adults were more highly tolerant of UVR than nauplii. Late-instar larvae of the predatory insect Chaoborus were more UVR tolerant than earlier instars. UVR tolerance showed no relationship to the UVR transparency of the source lake. Differential UVR tolerances among zooplankton taxa may alter community and ecosystem structure and function during anticipated changes in underwater UVR environments.

Climate change disrupts ecological systems in many ways. Many documented responses depend on species' life histories, contributing to the view that climate change effects are important but difficult to characterize generally. However, systematic variation in metabolic effects of temperature across trophic levels suggests that warming may lead to predictable shifts in food web structure and productivity. We experimentally tested the effects of warming on food web structure and productivity under two resource supply scenarios. Consistent with predictions based on universal metabolic responses to temperature, we found that warming strengthened consumer control of primary production when resources were augmented. Warming shifted food web structure and reduced total biomass despite increases in primary productivity in a marine food web. In contrast, at lower resource levels, food web production was constrained at all temperatures. These results demonstrate that small temperature changes could dramatically shift food web dynamics and provide a general, species-independent mechanism for ecological response to environmental temperature change.

Biological receptors used to detect underwater light vary in structure, from simple clusters of cells that only detect light intensity to complex organs that form detailed images. While most aquatic organisms perceive some portion of the visible spectrum, UV photoreceptors have recently been found in a wide diverse of species, from bacteria to fish. These UV photoreceptors are thought to aid in communication, foraging, and avoidance of damaging sunlight. However, they are not without costs as seeing in the UV can cause damage to the eye and image degradation. Given that the spectral composition of underwater light can directly affect an organism's physiology and behavior, environmental stressors that threaten to alter the color and clarity of fresh waters are a concern. These include nutrient and sediment loading associated with changes in climatic patterns and anthropogenic activities. This chapter discusses: (1) the underwater UV environment, (2) the structure and function of photoreceptors and the methods used to measure their spectral sensitivity, (3) the distribution of UV photoreceptors among freshwater organisms, (4) the adaptive significance of UV vision, and (5) potential effects of environmental stressors on UV vision.

[1] We generated suspended sediment solutions using river sediments and river water at concentrations similar to those observed during 1.5 year floods (Q1.5) and a dam removal (∼325 mg L−1) on the Deep River, North Carolina. Suspended sediment solutions were exposed to simulated solar radiation, equivalent to one clear, summer day at the study site (35°N). Concentrations of dissolved organic carbon (DOC), dissolved inorganic carbon (DIC), total dissolved nitrogen (TDN), dissolved inorganic nitrogen (DIN), dissolved organic nitrogen (DON), soluble reactive phosphorus (SRP) and total dissolved iron (Fed) were measured before and after exposure. Total dissolved carbon (TDC) budgets for each experiment were produced using DOC and DIC data. Sediment suspensions in the presence of simulated solar radiation were significant sources of dissolved C (119 ± 11 μmol C L−1 d−1; ± values indicate 1 standard error) and DON (1.7 ± 0.5 μmol N L−1 d−1) but not DIN or SRP. Extrapolations through the Deep River water column suggest that suspended sediments in the presence of light represent dissolved organic matter fluxes of 3.92 mmol C m−2 d−1 and 40 μmol N m−2 d−1. Additionally, sediment suspensions lowered river water Fed concentrations immediately (∼24%) and progressively (∼40–90%) in both light and dark treatments. Our research suggests suspended sediments in river ecosystems are potential sources of dissolved organic C and dissolved organic N while effectively removing Fed from the water column.

Seasonal increases in the ultraviolet (UV) transparency of the surface waters of an oligotrophic lake in Pennsylvania suggest that clear-water phase (CWP) events similar to those previously observed for visible light also exist for the potentially damaging UV wavelengths. Seasonal increases in zooplankton grazers indicate that they play a role in these changes in UV that is similar to the role that zooplankton play in CWP events involving longer-wavelength visible, or photosynthetically active radiation (PAR, 400¿700 nm). The potential role of zooplankton and incident UV in generating UV CWP events was investigated with a set of in situ microcosm experiments that manipulated UV and zooplankton, and followed changes in particulate and dissolved absorbance in the UV (320 nm) and PAR wavelength ranges over an 8-d period in April. Nutrients were also manipulated independently to examine the potential role of nutrient regeneration by zooplankton grazing in altering water transparency. Photobleaching by incident solar UV led to a strong and significant decrease in dissolved UV and PAR absorbance. The presence of zooplankton grazers also significantly decreased dissolved UV absorbance but increased dissolved PAR absorbance. Neither zooplankton nor UV had any significant effects on UV or PAR absorbance by particulates. In contrast, nutrient additions significantly increased dissolved absorbance in both the UV and PAR wavelength ranges, indicating that regeneration of nutrients by zooplankton offsets decreases in UV absorbance and enhances increases in PAR absorbance due to grazing. While photobleaching by UV radiation is likely to make a consistent strong contribution to UV CWP events in lakes, the net effects of zooplankton on UV transparency in a given lake will depend upon multiple factors including zooplankton density and a balance between the edibility and extent of nutrient limitation of the phytoplankton.

Phytoplankton biomass is commonly used as a water quality metric in the management of anthropogenic nitrogen and phosphorus loading, yet interpretation of the long-term response of phytoplankton biomass to nutrient regulation may be confounded by changes in zooplankton grazing pressure. Zooplankton community structure may be affected by planktivorous fish whose populations are subject to losses through fishing and gains through restoration efforts. We investigated temporal changes in phytoplankton biomass, nitrogen and phosphorus concentrations, zooplankton, and planktivorous fish in the tidal fresh and oligohaline Chowan River, NC, over a 30-year period in order to compare long-term trends in these parameters and infer their relative influence on phytoplankton biomass. Data were compiled from the records of two state agencies and several academic studies and supplemented by our own two year monitoring of water quality and zooplankton in the Chowan River mainstem and two tributaries. Seasonal trend decomposition using locally weighted regression was applied to chlorophyll a (a proxy for phytoplankton biomass), nitrogen, and phosphorus, and showed that chlorophyll a declined more quickly than did nutrient concentrations over the 30-year period. Despite the long-term decline in nitrogen and to a lesser extent phosphorus, the long-term trend in phytoplankton growth rate (predicted using an empirical model involving irradiance and nutrients) remained relatively constant. Zooplankton abundance increased from the period 1981–1982 to 2008–2010, as did the predicted zooplankton community water clearance rate (a proxy for zooplankton grazing). River herring, historically the dominant planktivore, declined dramatically over the 30-year period. The data indicate that the most parsimonious explanation for the long term decrease in chlorophyll a is an increase in zooplankton and not a decrease in nutrients. This inference leads us to hypothesize that the historically large river herring population exerted a positive, indirect influence on phytoplankton biomass in the Chowan River system, and that the decline in river herring has lead to higher zooplankton abundance and subsequently lower phytoplankton biomass.