In shallow water ecosytems the abiotic environment is more important than prey abundance for foraging terns

Shallow freshwater aquatic ecosystems are discrete feeding patches for fish eating birds. A unique feature of these ecosystems is that their physical conditions can change dramatically in a short period of time, particularly temperature, turbidity, and dissolved oxygen. Based on previous research we predicted that increasing turbidity will reduce the availability of fish to birds due to reduced visibility, while increasing temperature and decreasing dissolved oxygen will increase their availability through increases in activity and movement towards the more oxygenated surface areas, respectively. We also predicted that overall abundance of fish should increase feeding activity by terns. We measured these environmental variables, bird activity, and fish abundance from May to August from 2006 to 2008 in a marsh in southern Manitoba, Canada. Our results showed that only variation in dissolved oxygen levels affected feeding activity by terns. Since there was no relationship between bird and fish abundance either within or among years, these results suggest that it is the availability of prey (i.e. the upward movement of fish into the water column) and not their abundance per se that influences the number of avian predators present and hence the risk of predation to fish. These data demonstrate how the physical environment of aquatic ecosystems can impact terrestrial avian predators, and the link that exists between the physical environment and predator-prey interactions.     

Papyrus swamps and the respiratory ecology of Barbus neumayeri

Synopsis In seasonally flooding fresh waters, dissolved oxygen and fluctating water levels combine to create strong seasonal changes in habitat availability and dispersal pathways for fishes. In this study we demonstrate how respiratory mode can affect the use of and dispersal through hypoxic papyrus swamps for a small cyprinid fish, Barbus neumayeri, a species which uses aquatic surface respiration (ASR) in response to severe hypoxia. Monthly records of the distribution and relative abundance of B. neumayeri across 28 stations in a papyrus swamp in Uganda were used to examine seasonal patterns of habitat use and movement. The distribution of B. neumayeri was very restricted during the drier months. Most fish were captured in open pools and channels with large areas of open water surface and higher oxygen. Movement of B. neumayeri was limited to short periods during the wet season when peak water levels produced high levels of oxygen and pathways for movement. Three lines of evidence suggest that these patterns relate to its respiratory mode. These include: the positive :relationship between dissolved oxygen and the number of stations used per month, the positive relationship between fish density at dry season stations and dissolved oxygen, and the absence of fish from stations with. little open water surface area during the dry season. Significant differences in the gill morphology between B. neumayeri from the papyrus swamp and those from the main river suggest that papyrus swamps may contribute to the diversification of B. neumayeri by limiting movement and demanding specialization for extremely hypoxic waters.     

Parental Responses to Changes in Costs and Benefits Along an Environmental Gradient

We evaluated the effects of dissolved oxygen on offspring survival, parental costs, and the pattern of parental care in Florida flagfish, Jordanella floridae (Cyprinodontidae). Specifically, we quantified (1) embryonic development and survival in the absence of parental care, (2) behavior of non-reproductive adults, and (3) behavior of parental males across a gradient in dissolved oxygen. Embryo developmental rates and survivorship increased with dissolved oxygen, with a relatively sharp increase in survival between medium and high oxygen treatments. Non-reproductive adults increased their frequency of aquatic surface respiration, reduced overall activity, and increased opercular beat rate as oxygen declined, suggesting increased costs of activity with reduced oxygen. Taking these cost measures together, costs appear to increase slowly as oxygen starts to decline and then increase sharply as conditions approach hypoxia. In contrast, parental effort increased gradually with dissolved oxygen. We conclude that the increase in care from low to medium oxygen primarily results from a sharp decline in physiological costs, whereas the continued increase in care from medium to high oxygen primarily results from an increase in offspring value. In addition, our results highlight that the benefits of fanning for offspring are not well understood and that they may increase with oxygen, contrary to what has been previously assumed.     

Effects of hypoxia on the behavior and physiology of kelp forest fishes

Forecasts from climate models and oceanographic observations indicate increasing deoxygenation in the global oceans and an elevated frequency and intensity of hypoxic events in the coastal zone, which have the potential to affect marine biodiversity and fisheries. Exposure to low dissolved oxygen (DO) conditions may have deleterious effects on early life stages in fishes. This study aims to identify thresholds to hypoxia while testing behavioral and physiological responses of two congeneric species of kelp forest fish to four DO levels, ranging from normoxic to hypoxic (8.7, 6.0, 4.1, and 2.2 mg O₂/L). Behavioral tests identified changes in exploratory behavior and turning bias (lateralization), whereas physiological tests focused on determining changes in hypoxia tolerance (pCrit), ventilation rates, and metabolic rates, with impacts on the resulting capacity for aerobic activity. Our findings indicated that copper rockfish (Sebastes caurinus) and blue rockfish (Sebastes mystinus) express sensitivity to hypoxia; however, the strength of the response differed between species. Copper rockfish exhibited reduced absolute lateralization and increased escape time at the lowest DO levels, whereas behavioral metrics for blue rockfish did not vary with oxygen level. Both species exhibited decreases in aerobic scope (as a function of reduced maximum metabolic rate) and increases in ventilation rates to compensate for decreasing oxygen levels. Blue rockfish had a lower pCrit and stronger acclimation response compared to copper rockfish. The differences expressed by each species suggest that acclimatization to changing ocean conditions may vary, even among related species that recruit to the same kelp forest habitat, leading to winners and losers under future ocean conditions. Exposure to hypoxia can decrease individual physiological fitness through metabolic and aerobic depression and changes to anti‐predator behavior, with implications for the outcome of ecological interactions and the management of fish stocks in the face of climate change.     

Behavioral and physiological compensation for chronic hypoxia in the sailfin molly (Poecilia latipinna)

This study demonstrates that short-term behavioral and physiological responses may permit the sailfin molly Poecilia latipinna to cope successfully with extreme hypoxia and suggests an interaction between behavioral response (aquatic surface respiration [ASR]) and physiological compensation. Poecilia latipinna acclimated to chronic hypoxia (6 wk at 1.0 mg L(-1) O(2)) exhibited higher hemoglobin and red blood cell concentrations and a 17%-19% lower critical oxygen tension than fish acclimated to normoxia. Ventilation frequency increased twofold under acclimation to hypoxia, a response that did not diminish with time. However, the use of ASR was an immediate response to hypoxia that decreased over the acclimation period. This suggests that gradual physiological compensation decreases the threshold for ASR. There was no consistent effect of hypoxia on mortality and no effect of hypoxia treatment on the number of gestating females, suggesting that plastic behavioral and physiological responses in P. latipinna compensate for hypoxia to a degree that mitigates a decrease in survivorship and facilitates continued reproduction in a laboratory setting. However, there may be predation costs in the field related to ASR.     

Blood lactate and metabolic responses to controlled frequency breathing during graded swimming

Controlled frequency breathing (CFB) is a training technique used by swimmers in an effort to simulate high-intensity workloads by limiting oxygen availability to the body and stimulating anaerobic metabolism. During CFB, a swimmer voluntarily restricts breathing, which, theoretically, limits oxygen availability and stimulates anaerobic metabolism. The purpose of this study was to determine the influence of CFB on blood lactate and metabolic responses during graded increases in swimming intensity. A free swimming (FS) protocol was used to determine blood lactate and heart rate (HR) responses to CFB, while a tethered swimming (TS) protocol was used to determine blood lactate, HR, and ventilatory responses to CFB. The subjects swam four 3-minute trials at workloads of 55, 65, 75, and 85% of peak intensity during both protocols. A total of 46 competitive collegiate swimmers participated in the study. Thirty-four subjects (14 men and 20 women) completed the FS protocol, and 12 subjects (7 men and 5 women) completed the TS protocol. CFB reduced ventilation and Vo(2) (p < 0.05) during the TS protocol and reduced HR (p < 0.05) during the FS protocol when compared to normal breathing. However, CFB did not alter blood lactate concentrations for either protocol (p > 0.05). Our findings demonstrate that although CFB does not alter the blood lactate response to graded increases in swimming intensity, it appears to reduce the ventilatory and HR responses to exercise. Thus, swim coaches can use CFB at moderate intensities to simulate high-intensity training but should consider adjusting HR training zones to reflect the reduction in HR associated with reduced ventilation.     

Intestinal barrier function of Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis> L.) post smolts is reduced by common sea cage environments and suggested as a possible physiological welfare indicator

Background  

Fish farmed under high intensity aquaculture conditions are subjected to unnatural environments that may cause stress. Therefore awareness of how to maintain good health and welfare of farmed fish is important. For Atlantic salmon held in sea cages, water flow, dissolved oxygen (DO) levels and temperature will fluctuate over time and the fish can at times be exposed to detrimentally low DO levels and high temperatures. This experimental study investigates primary and secondary stress responses of Atlantic salmon post smolts to long-term exposure to reduced and fluctuating DO levels and high water temperatures, mimicking situations in the sea cages. Plasma cortisol levels and cortisol release to the water were assessed as indicators of the primary stress response and intestinal barrier integrity and physiological functions as indicators of secondary responses to changes in environmental conditions.     

Effects of hypoxia on movements and behavior of selected estuarine organisms from the southeastern United States

Hypoxia, or low dissolved oxygen, remains a common occurrence in estuarine waters as human activity in coastal areas expands. Fish kills, probably the most recognized indicator of these and other water quality problems, have significantly increased in recent years in many Southeastern United States estuaries. While entire aquatic communities are impacted by changes in available oxygen, estuarine organisms serve as appropriate indicators of these changes as they exhibit complex physiological and behavioral responses to hypoxia. The consequences of hypoxia for these species depend on their ability to detect and avoid areas of low dissolved oxygen. We conducted a series of two-way, replicated choice experiments with juvenile spot (Leiostomus xanthurus), pinfish (Lagodon rhomboides), croaker (Micropogonias undulatus), menhaden (Brevoortia tyrannus), white mullet (Mugil curema), mummichog (Fundulus heteroclitus), and brown shrimp (Penaeus aztecus) to determine their ability to detect and avoid specific levels of hypoxia. Additional data on organisms' movement patterns, aquatic surface respiration, and ventilation rates were collected. All species tested could detect and avoid 1 mgl(-1) dissolved oxygen. The hypoxia avoidance response differed among species, as some species exhibited an avoidance threshold while others exhibited a graded avoidance response. These data supply baseline information necessary to assess how some mobile estuarine organisms respond behaviorally to oxygen concentrations, and to understand how hypoxia more broadly impacts fish populations and estuarine community health.     

Temporal scale dependent interactions between multiple environmental disturbances in microcosm ecosystems

Global environmental change has negative impacts on ecological systems, impacting the stable provision of functions, goods, and services. Whereas effects of individual environmental changes (e.g. temperature change or change in resource availability) are reasonably well understood, we lack information about if and how multiple changes interact. We examined interactions among four types of environmental disturbance (temperature, nutrient ratio, carbon enrichment, and light) in a fully factorial design using a microbial aquatic ecosystem and observed responses of dissolved oxygen saturation at three temporal scales (resistance, resilience, and return time). We tested whether multiple disturbances combine in a dominant, additive, or interactive fashion, and compared the predictability of dissolved oxygen across scales. Carbon enrichment and shading reduced oxygen concentration in the short term (i.e. resistance); although no other effects or interactions were statistically significant, resistance decreased as the number of disturbances increased. In the medium term, only enrichment accelerated recovery, but none of the other effects (including interactions) were significant. In the long term, enrichment and shading lengthened return times, and we found significant two‐way synergistic interactions between disturbances. The best performing model (dominant, additive, or interactive) depended on the temporal scale of response. In the short term (i.e. for resistance), the dominance model predicted resistance of dissolved oxygen best, due to a large effect of carbon enrichment, whereas none of the models could predict the medium term (i.e. resilience). The long‐term response was best predicted by models including interactions among disturbances. Our results indicate the importance of accounting for the temporal scale of responses when researching the effects of environmental disturbances on ecosystems.     

Habitat degradation affects the summer activity of polar bears

Understanding behavioral responses of species to environmental change is critical to forecasting population-level effects. Although climate change is significantly impacting species’ distributions, few studies have examined associated changes in behavior. Polar bear (Ursus maritimus) subpopulations have varied in their near-term responses to sea ice decline. We examined behavioral responses of two adjacent subpopulations to changes in habitat availability during the annual sea ice minimum using activity data. Location and activity sensor data collected from 1989 to 2014 for 202 adult female polar bears in the Southern Beaufort Sea (SB) and Chukchi Sea (CS) subpopulations were used to compare activity in three habitat types varying in prey availability: (1) land; (2) ice over shallow, biologically productive waters; and (3) ice over deeper, less productive waters. Bears varied activity across and within habitats with the highest activity at 50–75% sea ice concentration over shallow waters. On land, SB bears exhibited variable but relatively high activity associated with the use of subsistence-harvested bowhead whale carcasses, whereas CS bears exhibited low activity consistent with minimal feeding. Both subpopulations had fewer observations in their preferred shallow-water sea ice habitats in recent years, corresponding with declines in availability of this substrate. The substantially higher use of marginal habitats by SB bears is an additional mechanism potentially explaining why this subpopulation has experienced negative effects of sea ice loss compared to the still-productive CS subpopulation. Variability in activity among, and within, habitats suggests that bears alter their behavior in response to habitat conditions, presumably in an attempt to balance prey availability with energy costs.     

Effects of predators and food availability on activity and growth of Chironomus tentans (Chironomidae: Diptera)

1. Larvae of Chironomus tentans Fab, decreased the amount of time they spent outside their tubes as the presence of predatory pumpkinseed sunfish (Lepomis gibbosus L.) increased. Greatest reductions in activity occurred at low levels of fish presence; above a certain level further increases in fish presence had little effect on activity. 2. Whether the pattern of predator presence was ordered or random had no effect on larval behaviour. Larvae did not habituate to short- or long-term predator presence. 3. Larvae were less active when more food was available and predator-induced reductions in activity were negatively related to food availability. Larval activity was much higher in the dark than it was in the light. 4. Over 7 days, presence of fish reduced the proportion of third-instar larvae that moulted but did not affect head width or dry mass; low food availability reduced the number of larvae that moulted as well as head width and dry mass of larvae in the fourth instar. 5. Results indicate that the behavioural response of larval chironomids to predator presence depends strongly on environmental conditions and that estimating the developmental costs of these behavioural responses under field conditions will be complicated.     

Potential impacts of global climate change on freshwater fisheries

Despite uncertainty in all levels of analysis, recent and long-term changes in our climate point to the distinct possibility that greenhouse gas emissions have altered mean annual temperatures, precipitation and weather patterns. Modeling efforts that use doubled atmospheric CO₂ scenarios predict a 1–7°C mean global temperature increase, regional changes in precipitation patterns and storm tracks, and the possibility of “surprises” or sudden irreversible regime shifts. The general effects of climate change on freshwater systems will likely be increased water temperatures, decreased dissolved oxygen levels, and the increased toxicity of pollutants. In lotic systems, altered hydrologic regimes and increased groundwater temperatures could affect the quality of fish habitat. In lentic systems, eutrophication may be exacerbated or offset, and stratification will likely become more pronounced and stronger. This could alter food webs and change habitat availability and quality. Fish physiology is inextricably linked to temperature, and fish have evolved to cope with specific hydrologic regimes and habitat niches. Therefore, their physiology and life histories will be affected by alterations induced by climate change. Fish communities may change as range shifts will likely occur on a species level, not a community level; this will add novel biotic pressures to aquatic communities. Genetic change is also possible and is the only biological option for fish that are unable to migrate or acclimate. Endemic species, species in fragmented habitats, or those in east–west oriented systems will be less able to follow changing thermal isolines over time. Artisanal, commercial, and recreational fisheries worldwide depend upon freshwater fishes. Impacted fisheries may make it difficult for developing countries to meet their food demand, and developed countries may experience economic losses. As it strengthens over time, global climate change will become a more powerful stressor for fish living in natural or artificial systems. Furthermore, human response to climate change (e.g., increased water diversion) will exacerbate its already-detrimental effects. Model predictions indicate that global climate change will continue even if greenhouse gas emissions decrease or cease. Therefore, proactive management strategies such as removing other stressors from natural systems will be necessary to sustain our freshwater fisheries.     

Effects of season and body-size on the distribution and diet of the Iberian chub Leuciscus pyrenaicus in a lowland catchment

Seasonal and ontogenetic variations in space and food use by Leuciscus pyrenaicus were analysed in a Portuguese lowland catchment. Large fish occurred mostly in deep permanent-flowing sites whereas small fish occurred mostly in the shallowest intermittent-flowing site. No seasonal or size-related changes in feeding intensity were found, but the diet changed both across seasons and throughout ontogeny. The diet was dominated by aquatic prey over all seasons, but during winter and summer more plant material and terrestrial prey, respectively, were eaten. Throughout ontogeny fish shifted from soft-bodied to hard-shelled prey and decreased animal prey breadth. Mean prey size increased with fish size but the prey size spectrum was more variable for medium-sized fish than for either small or large fish. It is suggested that: (i) large fish avoid shallow drying areas owing to the risk of mortality, either by thermal and respiratory stresses or increased predation by mammals and birds; (ii) seasonal changes in diet are a response to differences in prey availability; and (iii) morphological constraints, prey handling costs and habitat partitioning are responsible for size-related changes in diet.     

Interaction of a biotic factor (predator presence) and an abiotic factor (low oxygen) as an influence on benthic invertebrate communities

We examined the response of benthic invertebrates to hypoxia and predation risk in bioassay and behavioral experiments. In the bioassay, four invertebrate species differed widely in their tolerance of hypoxia. The mayfly, Callibaetis montanus, and the beetle larva, Hydaticus modestus, exhibited a low tolerance of hypoxia, the amphipod, Gammarus lacustris, was intermediate in its response and the caddisfly, Hesperophylax occidentalis, showed high tolerance of hypoxia. In the behavioral experiments, we observed the response of these benthic invertebrates, which differ in locomotor abilities, to vertical oxygen and temperature gradients similar to those in an ice-covered pond. With adequate oxygen, invertebrates typically remained on the bottom substrate. As benthic oxygen declined in the absence of fish, all taxa moved above the benthic refuge to areas with higher oxygen concentrations. In the presence of fish mayflies increased activity whereas all other taxa decreased activity in response to hypoxia. Mayflies and amphipods remained in the benthic refuge longer and endured lower oxygen concentrations whereas the vertical distribution of caddisflies and beetle larvae was not influenced by the presence of fish. As benthic oxygen declined in the presence of fish, all but the beetle larva reduced activity over all oxygen concentrations compared to when fish were absent. As benthic oxygen continued to decline, mayflies and amphipods moved above the benthic refuge and were preyed upon by fish. Thus, highly mobile taxa unable to tolerate hypoxia (mayflies and amphipods) responded behaviorally to declining oxygen concentrations by migrating upward in the water column. Taxa that were less mobile (beetle larvae) or hypoxia-tolerant (caddisflies) showed less of a response. Taxa most vulnerable to fish predation (mayflies and amphipods) showed a stronger behavioral response to predator presence than those less vulnerable (caddisflies and beetle larvae). Because invertebrates differ in their ability to withstand hypoxia, episodes of winter hypoxia could have long-lasting effects on benthic invertebrate communities either by direct mortality or selective predation on less tolerant taxa.     

The role of air breathing in the resistance of bimodally respiring fish to waterborne toxins

The bimodally respiring catfish Clarias macrocephalus Günther responded to a toxic extract of Croton tiglium (Euphorbiaceae) seeds by increased air breathing under both normoxic (8.1 ± 0.4 mgO₂ l⁻¹) and hypoxic (0.7 ± 0.1 mgO₂l⁻¹) conditions. Fish in hypoxia survived longer than those in normoxia when surface access was provided. When air breathing was prevented, survival time in toxin was greatly reduced at both levels of dissolved oxygen, and fish in normoxia survived longer than those in hypoxia. Non-toxin controls without surface access survived in normoxia but in hypoxia died at the same time as the fish in toxin. These results suggest that air breathing increases the resistance offish to toxins by permitting a decrease in the rate of gill ventilation and hence the rate at which toxins are absorbed.     

Effects of weather‐related episodic events in lakes: an analysis based on high‐frequency data

1. Weather‐related episodic events are typically unpredictable, and their duration is often short. Abiotic and biological responses are often missed in routine monitoring. These responses are, however, now of particular relevance given projected changes in extreme weather conditions. 2. We present data from high‐frequency monitoring stations from lakes in Europe, North America and Asia that illustrate two classes of abiotic effects of weather events: (i) generally short‐lived effects of storms on lake thermal structure and (ii) the more prolonged effects of high rainfall events on dissolved organic matter levels and water clarity. We further relate these abiotic effects to changes in dissolved oxygen or in chlorophyll a levels. 3. Three differing causes for weather‐related decreases in surface dissolved oxygen levels were observed: (i) entrainment of anoxic water from depth, (ii) reduction in primary productivity and (iii) increased mineralisation of organic carbon delivered from the catchment. 4. The duration of in‐lake effects tended to be longer for events driven by weather conditions with a longer return period, that is, conditions that were relatively more severe and less frequent at a site. While the susceptibility of lakes to change was related in part to the severity of the meteorological drivers, the impacts also depended on site‐specific factors in some cases. 5. The availability of high‐frequency data at these sites provided insight into the capacity of the lakes to absorb current and future pressures. Several of the changes we observed, including increases in carbon availability, decreases in photosynthetically active radiation and increased disturbance, have the capacity to shift lakes towards an increased degree of heterotrophy. The magnitude and direction of any such change will, however, also depend on the magnitude and direction of climate change for a given location and on lake and catchment characteristics.     

Photoperiodic regulation of the orexigenic effects of ghrelin in Siberian hamsters

Animals living in temperate climates with predictable seasonal changes in food availability may use seasonal information to engage different metabolic strategies. Siberian hamsters decrease costs of thermoregulation during winter by reducing food intake and body mass in response to decreasing or short-day lengths (SD). These experiments examined whether SD reduction in food intake in hamsters is driven, at least in part, by altered behavioral responses to ghrelin, a gut-derived orexigenic peptide which induces food intake via NPY-dependent mechanisms. Relative to hamsters housed in long-day (LD) photoperiods, SD hamsters consumed less food in response to i.p. treatment with ghrelin across a range of doses from 0.03 to 3 mg/kg. To determine whether changes in photoperiod alter behavioral responses to ghrelin-induced activation of NPY neurons, c-Fos and NPY expression were quantified in the arcuate nucleus (ARC) via double-label fluorescent immunocytochemistry following i.p. treatment with 0.3 mg/kg ghrelin or saline. Ghrelin induced c-Fos immunoreactivity (-ir) in a greater proportion of NPY-ir neurons of LD relative to SD hamsters. In addition, following ghrelin treatment, a greater proportion of ARC c-Fos-ir neurons were identifiable as NPY-ir in LD relative to SD hamsters. Changes in day length markedly alter the behavioral response to ghrelin. The data also identify photoperiod-induced changes in the ability of ghrelin to activate ARC NPY neurons as a possible mechanism by which changes in day length alter food intake.     

Evaluation of an alternative to feeding whole frozen fish in belugas (Delphinapterus leucas)

Feeding fish to captive piscivores can be challenging owing to cost, availability, variability in nutrient, and caloric composition, as well as handling and storage concerns. This trial evaluated the response of three belugas to being fed Fish Analog, an alternative to frozen fish. Body condition, gut transit time, serum chemistry and metabolic hormone analytes, immune function, and behavioral motivation were the dependent variables. Belugas (n=3) were fed various levels of Fish Analog (0-50%) over a 6-month period, and follow-up studies were conducted to further examine several dependent variables. When provided in gradually increasing amounts, belugas consumed the Fish Analog, with only minor fecal consistency changes and without behavioral responses indicative of gastric discomfort. Axillary girth and blubber thickness were positively correlated, and did not differ significantly with changes in the percentage of Fish Analog fed. Individual animal variation in initial passage time, some serum chemistry analytes, and immune function differences were noted following feeding of Fish Analog. Feeding Fish Analog reduced blood n9 fatty acids compared with captive belugas fed no Fish Analog. Feeding a DHA-enriched Fish Analog increased several n3 fatty acids, including eicosapentaenoic acid, but not DHA, compared with whales fed no Fish Analog or non-DHA-enriched Fish Analog. Fish Analog was shown to be a viable alternative to feeding fish at up to 50% of the dietary caloric density.     

Does browning-induced light limitation reduce fish body growth through shifts in prey composition or reduced foraging rates?

1. Browning of waters, coupled to climate change and land use changes, can strongly affect aquatic ecosystems. Browning-induced light limitation may have negative effects on aquatic consumers via shifts in resource composition and availability and by negatively affecting foraging of consumers relying on vision. However, the extent to which light limitation caused by browning affects fish via either of these two pathways is largely unknown.
2. Here we specifically test if fish growth responses to browning in a pelagic food web are best explained by changes in resource availability and composition due to light limitation, or by reduced foraging rates due to decreased visual conditions.
3. To address this question, we set up a mesocosm experiment to study growth responses of two different fish species to browning and conducted an aquaria experiment to study species-specific fish foraging responses to browning. Furthermore, we used a space-for-time approach to analyse fish body length-at-age across >40 lakes with a large gradient in lake water colour to validate experimental findings on species-specific fish growth responses.
4. With browning, we found an increase in chlorophyll a concentrations, shifts in zooplankton community composition, and a decrease in perch (Perca fluviatilis) but not roach (Rutilus rutilus) body growth. We conclude that fish growth responses are most likely to be linked to the observed shift in prey (zooplankton) composition. In contrast, we found limited evidence for reduced perch, but not roach, foraging rates in response to browning. This suggests that light limitation led to lower body growth of perch in brown waters mainly through shifts in resource composition and availability, perhaps in combination with decreased visibility. Finally, with the lake study we confirmed that perch but not roach body growth and length-at-age are negatively affected by brown waters in the wild.
5. In conclusion, using a combination of experimental and observational data, we show that browning of lakes is likely to (continue to) result in reductions in fish body growth of perch, but not roach, as a consequence of shifts in prey availability and composition, and perhaps reduced foraging.