Light pollution of freshwater ecosystems: principles, ecological impacts and remedies.

Light pollution caused by artificial light at night (ALAN) is increasingly recognized as a major driver of global environmental change. Since emissions are rapidly growing in an urbanizing world and half of the human population lives close to a freshwater shoreline, rivers and lakes are ever more exposed to light pollution worldwide. However, although light conditions are critical to aquatic species, and freshwaters are biodiversity hotspots and vital to human well-being, only a small fraction of studies conducted on ALAN focus on these ecosystems. The effects of light pollution on freshwaters are broad and concern all levels of biodiversity. Experiments have demonstrated diverse behavioural and physiological responses of species, even at low light levels. Prominent examples are skyglow effects on diel vertical migration of zooplankton and the suppression of melatonin production in fish. However, responses vary widely among taxa, suggesting consequences for species distribution patterns, potential to create novel communities across ecosystem boundaries, and cascading effects on ecosystem functioning. Understanding, predicting and alleviating the ecological impacts of light pollution on freshwaters requires a solid consideration of the physical properties of light propagating in water and a multitude of biological responses. This knowledge is urgently needed to develop innovative lighting concepts, mitigation strategies and specifically targeted measures. This article is part of the theme issue 'Light pollution in complex ecological systems'.

A framework for untangling the consequences of artificial light at night on species interactions.

Although much evidence exists showing organismal consequences from artificial light at night (ALAN), large knowledge gaps remain regarding ALAN affecting species interactions. Species interactions occur via shared spatio-temporal niches among species, which may be determined by natural light levels. We review how ALAN is altering these spatio-temporal niches through expanding twilight or full Moon conditions and constricting nocturnal conditions as well as creating patches of bright and dark. We review literature from a database to determine if ALAN is affecting species interactions via spatio-temporal dynamics. The literature indicates a growing interest in ALAN and species interactions: 58% of the studies we analysed have been published since 2020. Seventy-five of 79 studies found ALAN altered species interactions. Enhancements and reductions of species interactions were equally documented. Many studies revealed ALAN affecting species interactions spatially, but few revealed temporal alterations. There are biases regarding species interactions and ALAN-most studies investigated predator-prey interactions with vertebrates as predators and invertebrates as prey. Following this literature review, we suggest avenues, such as remote sensing and animal tracking, that can guide future research on the consequences of ALAN on species interactions across spatial and temporal axes. This article is part of the theme issue 'Light pollution in complex ecological systems'.

Microcosm experiment combined with process-based modeling reveals differential response and adaptation of aquatic primary producers to warming and agricultural run-off.

Fertilizers, pesticides and global warming are threatening freshwater aquatic ecosystems. Most of these are shallow ponds or slow-flowing streams or ditches dominated by submerged macrophytes, periphyton or phytoplankton. Regime shifts between the dominance of these primary producers can occur along a gradient of nutrient loading, possibly triggered by specific disturbances influencing their competitive interactions. However, phytoplankton dominance is less desirable due to lower biodiversity and poorer ecosystem function and services. In this study, we combined a microcosm experiment with a process-based model to test three hypotheses: 1) agricultural run-off (ARO), consisting of nitrate and a mixture of organic pesticides and copper, differentially affects primary producers and enhances the risk of regime shifts, 2) warming increases the risk of an ARO-induced regime shift to phytoplankton dominance and 3) custom-tailored process-based models support mechanistic understanding of experimental results through scenario comparison. Experimentally exposing primary producers to a gradient of nitrate and pesticides at 22°C and 26°C supported the first two hypotheses. ARO had direct negative effects on macrophytes, while phytoplankton gained from warming and indirect effects of ARO like a reduction in the competitive pressure exerted by other groups. We used the process-based model to test eight different scenarios. The best qualitative fit between modeled and observed responses was reached only when taking community adaptation and organism acclimation into account. Our results highlight the importance of considering such processes when attempting to predict the effects of multiple stressors on natural ecosystems.

Facing the infinity: tackling large samples of challenging Chironomidae (Diptera) with an integrative approach.

Background: Integrative taxonomy is becoming ever more significant in biodiversity research as scientists are tackling increasingly taxonomically challenging groups. Implementing a combined approach not only guarantees more accurate species identification, but also helps overcome limitations that each method presents when applied on its own. In this study, we present one application of integrative taxonomy for the highly abundant and particularly diverse fly taxon Chironomidae (Diptera). Although non-biting midges are key organisms in merolimnic systems, they are often cast aside in ecological surveys because they are very challenging to identify and extremely abundant. Methods: Here, we demonstrate one way of applying integrative methods to tackle this highly diverse taxon. We present a three-level subsampling method to drastically reduce the workload of bulk sample processing, then apply morphological and molecular identification methods in parallel to evaluate species diversity and to examine inconsistencies across methods. Results: Our results suggest that using our subsampling approach, identifying less than 10% of a sample's contents can reliably detect >90% of its diversity. However, despite reducing the processing workload drastically, the performance of our taxonomist was affected by mistakes, caused by large amounts of material. We conducted misidentifications for 9% of vouchers, which may not have been recovered had we not applied a second identification method. On the other hand, we were able to provide species information in cases where molecular methods could not, which was the case for 14% of vouchers. Therefore, we conclude that when wanting to implement non-biting midges into ecological frameworks, it is imperative to use an integrative approach.

Hypotheses in urban ecology: building a common knowledge base.

Urban ecology is a rapidly growing research field that has to keep pace with the pressing need to tackle the sustainability crisis. As an inherently multi-disciplinary field with close ties to practitioners and administrators, research synthesis and knowledge transfer between those different stakeholders is crucial. Knowledge maps can enhance knowledge transfer and provide orientation to researchers as well as practitioners. A promising option for developing such knowledge maps is to create hypothesis networks, which structure existing hypotheses and aggregate them according to topics and research aims. Combining expert knowledge with information from the literature, we here identify 62 research hypotheses used in urban ecology and link them in such a network. Our network clusters hypotheses into four distinct themes: (i) Urban species traits & evolution, (ii) Urban biotic communities, (iii) Urban habitats and (iv) Urban ecosystems. We discuss the potentials and limitations of this approach. All information is openly provided as part of an extendable Wikidata project, and we invite researchers, practitioners and others interested in urban ecology to contribute additional hypotheses, as well as comment and add to the existing ones. The hypothesis network and Wikidata project form a first step towards a knowledge base for urban ecology, which can be expanded and curated to benefit both practitioners and researchers.

Warming lowers critical thresholds for multiple stressor-induced shifts between aquatic primary producers.

In aquatic ecosystems, excessive nutrient loading is a global problem that can induce regime shifts from macrophyte- to phytoplankton-dominated states with severe consequences for ecosystem functions. Most agricultural landscapes are sites of nutrient and pesticide loading, which can interact with other stressors (e.g., warming) in additive, antagonistic, synergistic or reversed forms. The effects of multiple stressors on the resilience of macrophyte-dominated states and on critical thresholds for regime shifts are, however, unknown. We test the effects of individual and combined stressors of warming, nitrate, and various pesticides typically found in agricultural run-off (ARO) on the growth of macrophytes, periphyton, and phytoplankton in microcosms. We applied a one-level replicated design to test whether ARO induces a regime shift and a multifactorial dose-response design to model stressor thresholds and disentangle stressor interactions along a gradient. The individual stressors did not induce a regime shift, but the full ARO did. Nitrate and pesticides acted synergistically, inducing a shift with increasing phytoplankton biomass and decreasing macrophyte biomass. Warming amplified this effect and lowered critical thresholds for regime shifts. Shallow aquatic ecosystems in agricultural landscapes affected by global warming thus increasingly risk shifting to a turbid, phytoplankton-dominated state, and negatively impacting ecosystem service provisioning. Multiple stressor interactions must be considered when defining safe operating spaces for aquatic systems.

Large-scale sampling of the freshwater microbiome suggests pollution-driven ecosystem changes.

Freshwater microbes play a crucial role in the global carbon cycle. Anthropogenic stressors that lead to changes in these microbial communities are likely to have profound consequences for freshwater ecosystems. Using field data from the coordinated sampling of 617 lakes, ponds, rivers, and streams by citizen scientists, we observed linkages between microbial community composition, light and chemical pollution, and greenhouse gas concentration. All sampled water bodies were net emitters of CO2, with higher concentrations in running waters, and increasing concentrations at higher latitudes. Light pollution occurred at 75% of sites, was higher in urban areas and along rivers, and had a measurable effect on the microbial alpha diversity. Genetic elements suggestive of chemical stress and antimicrobial resistances (IntI1, blaOX58) were found in 85% of sites, and were also more prevalent in urban streams and rivers. Light pollution and CO2 were significantly related to microbial community composition, with CO2 inversely related to microbial phototrophy. Results of synchronous nationwide sampling indicate that pollution-driven alterations to the freshwater microbiome lead to changes in CO2 production in natural waters and highlight the vulnerability of running waters to anthropogenic stressors.

The rising moon promotes mate finding in moths.

To counteract insect decline, it is essential to understand the underlying causes, especially for key pollinators such as nocturnal moths whose ability to orientate can easily be influenced by ambient light conditions. These comprise natural light sources as well as artificial light, but their specific relevance for moth orientation is still unknown. We investigated the influence of moonlight on the reproductive behavior of privet hawkmoths (Sphinx ligustri) at a relatively dark site where the Milky Way was visible while the horizon was illuminated by distant light sources and skyglow. We show that male moths use the moon for orientation and reach females significantly faster with increasing moon elevation. Furthermore, the choice of flight direction depended on the cardinal position of the moon but not on the illumination of the horizon caused by artificial light, indicating that the moon plays a key role in the orientation of male moths.

Multiple-stressor exposure of aquatic food webs: Nitrate and warming modulate the effect of pesticides.

Shallow lakes provide essential ecological and environmental services but are exposed to multiple stressors, including agricultural runoff (ARO) and climate warming, which may act on different target receptors disrupting their normal functioning. We performed a microcosm experiment to determine the individual and combined effects of three stressors-pesticides, nitrate and climate warming-on two trophic levels representative of communities found in shallow lakes. We used three submerged macrophyte species (Myriophyllum spicatum, Potamogeton perfoliatus, Elodea nuttallii), eight benthic or pelagic microalgal species and three primary consumer species (Daphnia magna, Lymnaea stagnalis, Dreissena polymorpha) with different feeding preferences for benthic and pelagic primary producers. Eight different treatments consisted of a control, only nitrate, a pesticide cocktail, and a combination of nitrate and pesticides representing ARO, each replicated at ambient temperature and +3.5°C, mimicking climate warming. Pesticides negatively affected all functional groups except phytoplankton, which increased. Warming and nitrate modified these effects. Strong but opposite pesticide and warming effects on Myriophyllum drove the response of the total macrophyte biomass. Nitrate significantly suppressed Myriophyllum final biomass, but not overall macrophyte and microalgal biomass. Nitrate and pesticides in combination caused a macrophyte decline, and the system tipped towards phytoplankton dominance. Strong synergistic or even reversed stressor interaction effects were observed for macrophytes or periphyton. We emphasize the need for more complex community- and ecosystem-level studies incorporating multiple stressor scenarios to define safe operating spaces.

A global agenda for advancing freshwater biodiversity research.

Global freshwater biodiversity is declining dramatically, and meeting the challenges of this crisis requires bold goals and the mobilisation of substantial resources. While the reasons are varied, investments in both research and conservation of freshwater biodiversity lag far behind those in the terrestrial and marine realms. Inspired by a global consultation, we identify 15 pressing priority needs, grouped into five research areas, in an effort to support informed stewardship of freshwater biodiversity. The proposed agenda aims to advance freshwater biodiversity research globally as a critical step in improving coordinated actions towards its sustainable management and conservation.

Microplastic inclusion in birch tree roots.

In this pilot study, microplastic beads (5-50 µm) were tagged with fluorescent dye and introduced to the soil of potted Betula pendula Roth. (silver birch) saplings during the growing season. After five months, root samples were examined using fluorescence- and confocal laser scanning microscopy. This paper presents the first documented indication of the incorporation of microplastic into root tissues of woody plants and discusses the phytoremediation potential of birch in soil with microplastic contamination.

Design and implementation of an illumination system to mimic skyglow at ecosystem level in a large-scale lake enclosure facility.

Light pollution is an environmental stressor of global extent that is growing exponentially in area and intensity. Artificial skyglow, a form of light pollution with large range, is hypothesized to have environmental impact at ecosystem level. However, testing the impact of skyglow at large scales and in a controlled fashion under in situ conditions has remained elusive so far. Here we present the first experimental setup to mimic skyglow at ecosystem level outdoors in an aquatic environment. Spatially diffuse and homogeneous surface illumination that is adjustable between 0.01 and 10 lx, resembling rural to urban skyglow levels, was achieved with white light-emitting diodes at a large-scale lake enclosure facility. The illumination system was enabled by optical modeling with Monte-Carlo raytracing and validated by measurements. Our method can be adapted to other outdoor and indoor skyglow experiments, urgently needed to understand the impact of skyglow on ecosystems.

Towards Insect-Friendly Road Lighting-A Transdisciplinary Multi-Stakeholder Approach Involving Citizen Scientists.

(1) The project "Tatort Streetlight" implements an insect-friendly road light design in a four year before-after, control-impact (BACI) approach involving citizen scientists. It will broaden the stakeholder interests from solely anthropogenic perspectives to include the welfare of insects and ecosystems. Motivated by the detrimental impacts of road lighting systems on insects, the project aims to find solutions to reduce the insect attraction and habitat fragmentation resulting from roadway illumination. (2) The citizen science approach invites stakeholders to take part and join forces for the development of a sustainable and environmentally friendly road lighting solution. Here, we describe the project strategy, stakeholder participation and motivation, and how the effects of the alternative road luminaire and lighting design can be evaluated. (3) The study compares the changes in (a) insect behavior, (b) night sky brightness, and (c) stakeholder participation and awareness. For this purpose, different experimental areas and stakeholders in four communities in Germany are identified. (4) The project transfers knowledge of adverse effects of improperly managed road illumination and interacts with various stakeholders to develop a new road lighting system that will consider the well-being of street users, local residents, and insects.

Impact of Different Wavelengths of Artificial Light at Night on Phototaxis in Aquatic Insects.

The use of artificial light at night (ALAN) is increasing exponentially worldwide and there is growing evidence that ALAN contributes to the decline of insect populations. One of the most conspicuous ecological effects is the strong attraction of ALAN to flying insects. In several studies, light sources with strong short-wavelength emissions have been shown to attract the highest numbers of flying insects. Furthermore, flying stages of aquatic insects are reported to be more vulnerable to ALAN than flying stages of terrestrial insects. This is concerning because freshwater habitats are likely affected by ALAN that originates from human activity centers, which are typically close to sources of freshwater. However, the effects of ALAN on aquatic insects, which spend their larval phase (amphibiotic insects) or their whole life cycle (fully aquatic insects) in freshwaters, are entirely understudied. Here, we investigated the phototaxis of aquatic insects to ALAN at different wavelengths and intensities. We used floating light traps and compared four, near-monochromatic, lights (blue, green, red, and yellow) at two different photopic light intensities in a ditch system, which was not exposed to ALAN previously. Similar to flying stages of (aquatic and terrestrial) insects, we found a strong positive phototaxis of aquatic life stages. However, in contrast to the flying stages, there is no clear preference for short-wavelength light. Overall, responsivity to wavelengths in the center of the visible range (green, yellow; 500-600 nm) was significant for all orders of aquatic insects studied, and the nymphs of Ephemeroptera did not respond to blue light at all. This is likely an adaption to how light is attenuated in freshwater systems, where not only the water itself but also a variety of optical constituents act as a color filter, often like in our case filtering out short-wavelength light. Therefore, insects living in freshwater bodies often live in longer wavelength-dominated environments and might therefore be especially sensitive to green/yellow light. In conclusion, the different spectral sensitivities of both aquatic and flying insects should be taken into account when planning lighting near freshwater.

Misbalance of thyroid hormones after two weeks of exposure to artificial light at night in Eurasian perch Perca fluviatilis.

Artificial light at night (ALAN) can affect the physiology and behavior of animals because it alters the natural rhythm of light and darkness. Thyroid hormones (TH) are partially regulated by the light information of photoperiod and are involved in metabolic adjustments to daily and seasonal changes in the environment, such as larval and juvenile development, somatic growth and reproduction. ALAN can change photoperiodic information and might thereby lead to changes in thyroid metabolism, but so far research on this topic is scarce. Therefore, we tested in two different experiments the effects of nocturnal illumination at a wide range of light intensities on TH in plasma of Eurasian perch (Perca fluviatilis). Total 3,3',5-triiodo-L-thyronine (T3) was significantly affected by ALAN and reduced at the highest tested intensity of 100 lx after only two weeks of exposure. Although total L-thyroxine (T4) was not significantly affected, the ratio of T3 to T4 tended to slightly decrease at 100 lx. In a second low-light experiment ALAN did not have clear effects on T3, T4 or the ratio of T3 to T4 at intensities between 0.01 lx and 1 lx. The results show first signs of endocrine disruption in thyroid metabolism after a relatively short ALAN exposure of two weeks under high-intensity streetlight conditions. Misbalanced thyroidal status can have serious implications for metabolic rates as well as developmental and reproductive processes.

Innate immunity, oxidative stress and body indices of Eurasian perch Perca fluviatilis after two weeks of exposure to artificial light at night.

Artificial light at night (ALAN) can disrupt biological rhythms of fish and other vertebrates by changing the light information of the nocturnal environment. Disrupted biorhythms can impair the immune system of vertebrates as it has been shown for conditions with continuous illumination or long-day photoperiod in many vertebrates, including fish. Nonetheless, this has not been shown so far for typical ALAN scenarios with high light intensities during day and low light intensities at night. Therefore, in this study, proxies for the innate immune system and oxidative stress as well as body indices of Eurasian perch Perca fluviatilis were measured under a wide range of intensities of nocturnal illumination. The authors found no changes in parameters of the innate immune system and no significant changes in proxies for oxidative stress after 2-week exposures to nocturnal illuminance ranging from 0.01 lx to 1 lx in one experiment or from 1 lx to 100 lx in a second experiment. A decrease in the hepato-somatic index at the highest tested light intensity of 100 lx compared to the dark control was the only significant difference in all parameters among treatments. After 2 weeks of exposure, ALAN does not seem to seriously challenge the innate immune system and seems to cause less oxidative stress than expected. The results of this study contradict the findings from other studies applying continuous illumination or long-day photoperiod and highlight the importance of further research in this field. Because ALAN represents a sustained modulation of the environment that may have cumulative effects over time, long-term studies are required for a better understanding of how ALAN modulates the health of fish.

Urban Lighting Research Transdisciplinary Framework-A Collaborative Process with Lighting Professionals.

Over the past decades, lighting professionals have influenced the experience of the night by brightly illuminating streets, buildings, skylines, and landscapes 24/7. When this became the accepted norm, a dual perspective on night-time was shaped and the visual enjoyment of visitors after dusk was prioritized over natural nightscapes (nocturnal landscapes). During this time, researchers of artificial light at night (ALAN) observed and reported a gradual increase in unnatural brightness and a shift in color of the night-time environment. As a consequence, ALAN has been identified as a relevant pollutant of aquatic and terrestrial habitats, and an environmental stressor, which may adversely affect a wide range of organisms, from micro-organisms to humans. Unfortunately, lighting professionals and ALAN researchers usually attempt to solve today's sustainable urban lighting problems distinctive to their fields of study, without a dialogue between research and practice. Therefore, in order to translate research knowledge as an applicable solution for the lighting practice and to minimize the impact on the environment, a collaborative framework involving a transdisciplinary process with lighting professionals is crucial to potentially bring the practice, research, production, decision-making, and planning closer to each other. This paper presents a framework to help reduce the existing gap of knowledge, because appropriate lighting applications depend upon it. Access to less light polluted nightscapes in urban environments is just as important as access to unpolluted water, food, and air. This call for action towards sustainable urban lighting should be included in future lighting policies to solve the urgent environmental and health challenges facing our world.

Light intensity and spectral distribution affect chytrid infection of cyanobacteria via modulation of host fitness.

Light gradients are an inherent feature in aquatic ecosystems and play a key role in shaping the biology of phytoplankton. Parasitism by chytrid fungi is gaining increasing attention as a major control agent of phytoplankton due to its previously overlooked ubiquity, and profound ecological and evolutionary consequences. Despite this interest, if and how light conditions modulate phytoplankton chytridiomycosis remains poorly studied. We investigated life-history traits of a chytrid parasite, Rhizophydium megarrhizum, under different light intensities and spectral compositions when infecting two closely related planktonic cyanobacteria with different light-harvesting strategies: Planktothrix rubescens and P. agardhii. In general, parasite transmission was highest under light conditions (both intensity and quality) that maximized growth rates for uninfected cyanobacteria. Chytrid encystment on hosts was significantly affected by light intensity and host strain identity. This likely resulted from higher irradiances stimulating the increased discharge of photosynthetic by-products, which drive parasite chemotaxis, and from strain-specific differences at the cell-surface. Comparisons of parasite transmission and host growth rates under different light conditions suggest the potential for epidemic development at higher irradiances, whereas host and parasite could coexist without epidemic outbreaks at lower light levels. These results illustrate the close relationship between parasite transmission and host fitness, which is ultimately modulated by the external environment.

Can skyglow reduce nocturnal melatonin concentrations in Eurasian perch?

Artificial light at night (ALAN) changes the natural rhythm of light and darkness and can impair the biorhythms of animals, for example the nocturnal melatonin production of vertebrates, which serves as a proxy for daily physiological rhythms. Freshwater fish are exposed to ALAN in large urban and suburban areas in the form of direct light or in the form of skyglow, a diffuse brightening of the night sky through the scattered light reflected by clouds, atmospheric molecules, and particles in the air. However, investigations on the sensitivity of melatonin production of fish towards low intensities of ALAN in the range of typical skyglow are rare. Therefore, we exposed Eurasian perch (Perca fluviatilis) to nocturnal illumination levels of 0.01 lx, 0.1 lx and 1 lx and a control group with dark nights and daylight intensities of 2900 lx in all groups. After ten days of exposure to the experimental conditions, tank water was non-invasively sampled every 3 h over a 24 h period and melatonin was measured by ELISA. Melatonin was gradually reduced in all treatments with increasing intensity of ALAN whereas rhythmicity was maintained in all treatment groups although at 1 lx not all evaluated parameters confirmed rhythmicity. These results show a high sensitivity of Eurasian perch towards ALAN indicating that low light intensities of 0.01 lx and 0.1 lx as they occur in urban and suburban areas in the form of skyglow can affect the physiology of Eurasian perch. Furthermore, we highlight how this may impact perch in their sensitivity towards lunar rhythms and the role of skyglow for biorhythms of temperate freshwater fish.