Shedding light with harmonic radar: Unveiling the hidden impacts of streetlights on moth flight behavior.

One of the most dramatic changes occurring on our planet is the ever-increasing extensive use of artificial light at night, which drastically altered the environment to which nocturnal animals are adapted. Such light pollution has been identified as a driver in the dramatic insect decline of the past years. One nocturnal species group experiencing marked declines are moths, which play a key role in food webs and ecosystem services such as plant pollination. Moths can be easily monitored within the illuminated area of a streetlight, where they typically exhibit disoriented behavior. Yet, little is known about their behavior beyond the illuminated area. Harmonic radar tracking enabled us to close this knowledge gap. We found a significant change in flight behavior beyond the illuminated area of a streetlight. A detailed analysis of the recorded trajectories revealed a barrier effect of streetlights on lappet moths whenever the moon was not available as a natural celestial cue. Furthermore, streetlights increased the tortuosity of flights for both hawk moths and lappet moths. Surprisingly, we had to reject our fundamental hypothesis that most individuals would fly toward a streetlight. Instead, this was true for only 4% of the tested individuals, indicating that the impact of light pollution might be more severe than assumed to date. Our results provide experimental evidence for the fragmentation of landscapes by streetlights and demonstrate that light pollution affects movement patterns of moths beyond what was previously assumed, potentially affecting their reproductive success and hampering a vital ecosystem service.

Species sensitivities to artificial light at night: A phylogenetically controlled multilevel meta-analysis on melatonin suppression.

The rapid urbanization of our world has led to a surge in artificial lighting at night (ALAN), with profound effects on wildlife. Previous research on wildlife's melatonin, a crucial mechanistic indicator and mediator, has yielded inconclusive evidence due to a lack of comparative analysis. We compiled and analysed an evidence base including 127 experiments with 437 observations across 31 wild vertebrates using phylogenetically controlled multilevel meta-analytic models. The evidence comes mainly from the effects of white light on melatonin suppression in birds and mammals. We show a 36% average decrease in melatonin secretion in response to ALAN across a diverse range of species. This effect was observed for central and peripheral melatonin, diurnal and nocturnal species, and captive and free-living populations. We also reveal intensity-, wavelength-, and timing-dependent patterns of ALAN effects. Exposure to ALAN led to a 23% rise in inter-individual variability in melatonin suppression, with important implications for natural selection in wild vertebrates, as some individuals may display higher tolerance to ALAN. The cross-species evidence has strong implications for conservation of wild populations that are subject to natural selection of ALAN. We recommend measures to mitigate harmful impacts of ALAN, such as using 'smart' lighting systems to tune the spectra to less harmful compositions.

Threatened chronotopes: can chronobiology help endangered species?

Pittendrigh and Daan's 1976 article "Pacemaker structure: A clock for all seasons" marks the foundation of modern seasonal chronobiology. It proposed the internal coincidence model comprised of a Morning (M) and Evening (E) oscillator, which are coupled but synchronized separately by dawn and dusk. It has become an attractive model to explain the seasonal adaptation of circadian rhythms. Using the example of the European hamster, this article connects the classical entrainment concept to species decline and, ultimately, conservation concepts. Seasonality of this species is well studied and circannual rhythms have been described in at least 32 parameters. The European hamster is listed as critically endangered on the International Union for Conservation of Nature (IUCN) red list. Changes in the temporal structure of the environment (the chronotope) caused by climate change and light pollution might be responsible for the global decline. The article shows that classical chronobiological concepts such as the internal coincidence model (Pittendrigh and Daan Pittendrigh and Daan, J Comp Physiol [a] 106:333-355, 1976) are helpful to understand the (chronobiological) causes of the decline and can potentially support species conservation. Knowing the species' physiological limitations as well as its adaptation capacities can potentially prevent its extinction at a time when classical conservation concepts have reached their limits.

Crickets in the spotlight: exploring the impact of light on circadian behavior.

Crickets serve as a well-established model organism in biological research spanning various fields, such as behavior, physiology, neurobiology, and ecology. Cricket circadian behavior was first reported over a century ago and prompted a wealth of studies delving into their chronobiology. Circadian rhythms have been described in relation to fundamental cricket behaviors, encompassing stridulation and locomotion, but also in hormonal secretion and gene expression. Here we review how changes in illumination patterns and light intensity differentially impact the different cricket behaviors as well as circadian gene expression. We further describe the cricket's circadian pacemaker. Ample anatomical manipulations support the location of a major circadian pacemaker in the cricket optic lobes and another in the central brain, possibly interconnected via signaling of the neuropeptide PDF. The cricket circadian machinery comprises a molecular cascade based on two major transcriptional/translational negative feedback loops, deviating somewhat from the canonical model of Drosophila and emphasizing the significance of exploring alternative models. Finally, the nocturnal nature of crickets has provided a unique avenue for investigating the repercussions of artificial light at night on cricket behavior and ecology, underscoring the critical role played by natural light cycles in synchronizing cricket behaviors and populations, further supporting the use of the cricket model in the study of the effects of light on insects. Some gaps in our knowledge and challenges for future studies are discussed.

Do seabirds dream of artificial lights? Understanding light preferences of Procellariiformes.

Seabirds, and particularly fledglings of burrow-nesting species, are greatly impacted by light pollution. During their inaugural flights from colony to sea, fledglings become grounded after encountering artificial light. Such groundings, or fallout events, affect many fledglings each year, causing mass mortality events. To mitigate this light-induced mortality, rescue programmes have been implemented for decades at many locations worldwide. Despite the notoriety of fallouts and their conservation implications, the contributing behavioural and biological factors remain mostly unknown. How the mechanisms of light attraction and light avoidance interact and how they manifest in different groups (e.g. age, personality, populations) or light pollution levels remain open questions. We tested behavioural choices of Cory's shearwater Calonectris borealis fledglings, rescued after being grounded in urban areas, and choices of breeding adults for contrasting light sources. Fledglings and adults were exposed to one of three treatments in an experimental Y-maze set-up: white light versus no light, blue versus red light, and a control with no light on each arm of the Y-maze. Both age groups clearly chose the no-light arms and the red light arm. This choice for longer wavelengths and darker environments, along with slower responses by fledglings, suggests that close range artificial light causes disorientation in seabirds. Our study helps to clarify the behavioural components of fallouts and provides further evidence on the disruptive effects of nocturnal artificial light on sensitive species like Procellariiformes.

Drivers of fatal bird collisions in an urban center.

Millions of nocturnally migrating birds die each year from collisions with built structures, especially brightly illuminated buildings and communication towers. Reducing this source of mortality requires knowledge of important behavioral, meteorological, and anthropogenic factors, yet we lack an understanding of the interacting roles of migration, artificial lighting, and weather conditions in causing fatal bird collisions. Using two decades of collision surveys and concurrent weather and migration measures, we model numbers of collisions occurring at a large urban building in Chicago. We find that the magnitude of nocturnal bird migration, building light output, and wind conditions are the most important predictors of fatal collisions. The greatest mortality occurred when the building was brightly lit during large nocturnal migration events and when winds concentrated birds along the Chicago lakeshore. We estimate that halving lighted window area decreases collision counts by 11× in spring and 6× in fall. Bird mortality could be reduced by ∼60% at this site by decreasing lighted window area to minimum levels historically recorded. Our study provides strong support for a relationship between nocturnal migration magnitude and urban bird mortality, mediated by light pollution and local atmospheric conditions. Although our research focuses on a single site, our findings have global implications for reducing or eliminating a critically important cause of bird mortality.

Long wavelength light reduces the negative consequences of dim light at night.

Many patients with autism spectrum disorders (ASD) show disturbances in their sleep/wake cycles, and they may be particularly vulnerable to the impact of circadian disruptors. We have previously shown that a 2-weeks exposure to dim light at night (DLaN) disrupts diurnal rhythms, increases repetitive behaviors and reduces social interactions in contactin-associated protein-like 2 knock out (Cntnap2 KO) mice. The deleterious effects of DLaN may be mediated by intrinsically photosensitive retinal ganglion cells (ipRGCs) expressing the photopigment melanopsin, which is maximally sensitive to blue light (480 nm). In this study, the usage of a light-emitting diode array enabled us to shift the spectral properties of the DLaN while keeping the intensity of the illumination at 10 lx. First, we confirmed that the short-wavelength enriched lighting produced strong acute suppression of locomotor activity (masking), robust light-induced phase shifts, and cFos expression in the suprachiasmatic nucleus in wild-type (WT) mice, while the long-wavelength enriched lighting evoked much weaker responses. Opn4DTA mice, lacking the melanopsin expressing ipRGCs, were resistant to DLaN effects. Importantly, shifting the DLaN stimulus to longer wavelengths mitigated the negative impact on the activity rhythms and 'autistic' behaviors (i.e. reciprocal social interactions, repetitive grooming) in the Cntnap2 KO as well as in WT mice. The short-, but not the long-wavelength enriched, DLaN triggered cFos expression in in the basolateral amygdala (BLA) as well as in the peri-habenula region raising that possibility that these cell populations may mediate the effects. Broadly, our findings are consistent with the recommendation that spectral properties of light at night should be considered to optimize health in neurotypical as well as vulnerable populations.

Dim light at night affects the locomotor activity of nocturnal African pygmy mice (Mus minutoides) in an intensity-dependent manner.

Rodents are integral components of ecosystems as they provide several important ecosystem services. Despite their importance as prey, pollinators and seed distributors, African rodents are largely understudied. The effect of anthropogenic changes such as artificial light at night extends past urban areas to peri-urban and rural habitats, and can have profound effects on entire ecosystems. We investigated the effect of dim light at night (dLAN) on the locomotor activity rhythms of the African pygmy mouse (Mus minutoides). Pygmy mice showed a dramatic, intensity-dependent reduction in their locomotor activity when subjected to dLAN, which was accompanied by a delay in the activity onset. We also considered masking responses with a dark pulse (DP) during the day and a light pulse at night. All animals became inactive in response to a light pulse during the night, whereas approximately half of the animals showed activity during a DP in the day. Our results suggest that the African pygmy mouse is highly sensitive to light and that their activity is strongly masked by light. In their natural environment, vegetation could shield pygmy mice against high light levels; however, other anthropogenic disturbances can alter the behaviour of these animals and could affect their survival.

Evaluating the effects of moonlight on the vertical flight profiles of three western palaearctic swifts.

Recent studies have suggested the presence of moonlight mediated behaviour in avian aerial insectivores, such as swifts. Here, we use the combined analysis of state-of-the-art activity logger data across three swift species, the common, pallid and alpine swifts, to quantify flight height and activity in responses to moonlight-driven crepuscular and nocturnal light conditions. Our results show a significant response in flight heights to moonlight illuminance for common and pallid swifts, i.e. when moon illuminance increased flight height also increased, while a moonlight-driven response is absent in alpine swifts. We show a weak relationship between night-time illuminance-driven responses and twilight ascending behaviour, suggesting a decoupling of both crepuscular and night-time behaviour. We suggest that swifts optimize their flight behaviour to adapt to favourable night-time light conditions, driven by light-responsive and size-dependent vertical insect stratification and weather conditions.

Light pollution enhances ground-level exposure to airborne toxic chemicals for nocturnally migrating passerines.

Anthropogenic activities generate different forms of environmental pollution, including artificial light at night (ALAN) and airborne toxic chemicals (ATCs). Nocturnally migrating birds are attracted to ALAN during migration and if ALAN occurs in unison with ATC, the chances of ground-level ATC contamination occurring at stopover sites could increase. Here, we document the relationship between ALAN and ATC within the contiguous United States based on 479 toxic chemicals from 15,743 releasing facilities. Using weekly diurnal estimates of relative abundance for 165 nocturnally migrating passerine (NMP) bird species, we assess how the species richness and relative abundance of NMP species are correlated with ALAN and ATC across the annual cycle. The concentration of ATC increased with increasing ALAN levels, except at the highest ALAN levels. The species richness of NMP species was positively correlated with ATC during the non-breeding season and migration, and negatively correlated during the breeding season. The relative abundance of NMP species was negatively correlated with ATC during the breeding and non-breeding seasons and the correlation did not differ from zero during migration. Through the disorienting influence of ALAN, our findings suggest large numbers of NMP species are being exposed to higher ATC concentrations at stopover sites. Outside of migration, large numbers of NMP species that winter along the US Gulf Coast are being exposed for an extended period of time to higher ATC concentrations. Initiatives designed to decrease ALAN during migration have the potential to reduce the acute and chronic effects of ATC contamination, lower the maternal transfer of toxic chemicals to eggs, and decrease the biologically mediated transport of toxic chemicals across regions. However, these initiatives will not benefit species that experience prolonged ATC exposure during the non-breeding season along the US Gulf Coast, a region that could be a significant source of ATC contamination for North American birds.

Stress tolerance is influenced by artificial light at night during development and life-history strategy.

Artificial light at night (ALAN) is increasingly prevalent worldwide, but life-history strategy may mitigate the costs of ALAN for animals. Yet, interactions among ALAN, life-history strategy and tolerance to climate-related stressors are unknown. We determined if developmental ALAN exposure (1) affects development, (2) affects adult phenotype, including heat and desiccation tolerance, and (3) affects and/or interacts with life-history strategy. We used the variable field cricket (Gryllus lineaticeps) because its geographic range is increasingly exposed to ALAN, heat, and drought conditions, and it exhibits different life-history strategies (flight-capability versus flight-incapability). ALAN affected adult phenotype, with positive effects on body mass (and size) and female reproductive investment, and a negative effect on heat tolerance. Life-history strategy also affected stress tolerance; flight-incapable females had greater heat tolerance and their desiccation tolerance was improved by ALAN exposure. Key features of environmental change (i.e. exposure to ALAN, heat and drought) may favor some life-history strategies over others.

OSpRad: an open-source, low-cost, high-sensitivity spectroradiometer.

Spectroradiometry is a vital tool in a wide range of biological, physical, astronomical and medical fields, yet its cost and accessibility are frequent barriers to use. Research into the effects of artificial light at night (ALAN) further compounds these difficulties with requirements for sensitivity to extremely low light levels across the ultraviolet to human-visible spectrum. Here, I present an open-source spectroradiometry (OSpRad) system that meets these design challenges. The system utilises an affordable miniature spectrometer chip (Hamamatsu C12880MA), combined with an automated shutter and cosine-corrector, microprocessor controller, and graphical user interface 'app' that can be used with smartphones or desktop computers. The system has high ultraviolet sensitivity and can measure spectral radiance at 0.001 cd m-2 and irradiance at 0.005 lx, covering the vast majority of real-world night-time light levels. The OSpRad system's low cost and high sensitivity make it well suited to a range of spectrometry and ALAN research.

Artificial light impairs local attraction to females in male glow-worms.

The negative effects of artificial lighting at night (ALAN) on insects are increasingly recognised and have been postulated as one possible cause of declines in insect populations. Yet, the behavioural mechanisms underpinning ALAN effects on insects remain unclear. ALAN interferes with the bioluminescent signal female glow-worms use to attract males, disrupting reproduction. To determine the behavioural mechanisms that underpin this effect of ALAN, we quantified the effect of white illumination on males' ability to reach a female-mimicking LED within a Y-maze. We show that as the intensity of illumination increases, the proportion of males reaching the female-mimicking LED declines. Brighter illumination also increases the time taken by males to reach the female-mimicking LED. This is a consequence of males spending more time: (i) in the central arm of the Y-maze; and (ii) with their head retracted beneath their head shield. These effects reverse rapidly when illumination is removed, suggesting that male glow-worms are averse to white light. Our results show that ALAN not only prevents male glow-worms from reaching females, but also increases the time they take to reach females and the time they spend avoiding exposure to light. This demonstrates that the impacts of ALAN on male glow-worms extend beyond those previously observed in field experiments, and raises the possibility that ALAN has similar behavioural impacts on other insect species that remain undetected in field experiments.

Experimental light at night explains differences in activity onset between urban and forest great tits.

Artificial light at night (ALAN) is rapidly increasing and so is scientific interest in its ecological and evolutionary consequences. In wild species, ALAN can modify and disrupt biological rhythms. However, experimental proof of such effects of ALAN in the wild is still scarce. Here, we compared diel rhythms of incubation behaviour, inferred from temperature sensors, of female great tits (Parus major) breeding in urban and forest sites. In parallel, we simulated ALAN by mounting LED lights (1.8 lx) inside forest nest-boxes, to determine the potentially causal role of ALAN affecting diel patterns of incubation. Urban females had an earlier onset of activity compared to forest females. Experimentally ALAN-exposed forest females were similar to urban females in their advanced onset of activity, compared to unexposed forest birds. However, forest females exposed to experimental ALAN, but not urban females, were more restless at night than forest control females. Our findings demonstrate that ALAN can explain the early activity timing in incubating urban great tits, but its effects on sleep disturbance in the forest are not reflected in urban females. Consequently, future research needs to address potential effects of ALAN-induced timing on individual health, fitness and population dynamics, in particular in populations that were not previously affected by light pollution.

Outdoor artificial light at night and human health: A review of epidemiological studies.

We conducted a non-systematic review of epidemiological studies on a potential link between exposure to outdoor artificial light at night (O-ALAN) and disease occurrence in humans published since 2009. In recent years, a number of presses have been published on this issue, but the conclusions have been mixed. We therefore decided to critically analyze the available epidemiological evidence of such a correlation. After a careful search, 51 studies were identified and included in the review. They addressed the potential link between O-ALAN exposure and the incidence of breast cancer, other cancers, sleep and circadian rhythm disorders, obesity and cardiovascular diseases, mental disorders, infectious diseases, and complications during pregnancy and childbirth. The vast majority of papers revealed the existence of such a link. However, the amount of epidemiological evidence supporting the correlation across groups of disorders varied widely. In addition, we found that all papers contained at least one of the following omissions: lack of the temporal and spatial resolution in light at night measurements, measuring only light intensity without considering its wavelength, and not accounting for many important confounding factors in their statistical analyses. Therefore, we believe that the link between O-ALAN exposure and the occurrence of the disorders in question suggested by the authors of the reviewed papers may be in some cases at least to some extent, a coincidence. Further epidemiological studies, free of significant omissions highlighted in this paper, are needed.

Impact of the exposome on cardiovascular disease.

Air pollution, noise pollution, and light pollution have emerged as important but often overlooked risk factors for cardiovascular disease. In this review, we examine the emerging concept of the exposome, highlighting the close relationship between environmental exposure (e.g. PM2.5, traffic noise, and night light) and cardiovascular disease, finally addressing the possible mitigation strategies that should be implemented to reduce the impact of air, noise, and light pollution on cardiovascular morbidity and mortality.

Artifical light at night triggers slight transcriptomic effects on melatonin signaling but not synthesis in tadpoles of two anuran species.

The worldwide expansion of artificial light at night (ALAN) is acknowledged as a threat to biodiversity through alterations of the natural photoperiod triggering the disruption of physiological functions. In vertebrates, melatonin production during the dark phase can be decreased or suppressed by nocturnal light as shown in many taxa. But the effect of ALAN at low intensity mimicking light pollution in peri-urban area has never been investigated in amphibians. We filled this gap by studying the impact of low ALAN levels on the expression of genes related to melatonin synthesis and signaling in two anurans (agile frog, Rana dalmatina, and common toad, Bufo bufo). Circadian expression of genes encoding enzymes catalyzing melatonin synthesis (aralkylamine N-acetyltransferase, AANAT and acetylserotonin O-methyltransferase, ASMT) or melatonin receptors (Mel1a, Mel1b and Mel1c) was investigated using RT-qPCR after 23 days of nocturnal exposure to control (< 0.01 lx) or low ALAN (3 lx). We showed that the relative abundance of most transcripts was low in late afternoon and early evening (06 pm and 08 pm) and increased throughout the night in R. dalmatina. However, a clear and ample nocturnal pattern of target gene expression was not detected in control tadpoles of both species. Surprisingly, a low ALAN level had little influence on the relative expression of most melatonin-related genes. Only Mel1c expression in R. dalmatina and Mel1b expression in B. bufo were affected by ALAN. This target gene approach provides experimental evidence that melatonin signaling pathway was slightly affected by low ALAN level in anuran tadpoles.

Insulin ameliorates dim blue light at night-induced apoptosis in hippocampal neurons via the IR/IRS1/AKT/GSK3ß/ß-catenin signaling pathway.

In recent years, the damaging effects of night light pollution, one of the environmental pollutions, on memory has been attracting attention. However, the underlying molecular mechanisms by which light at night, especially blue light at night, impairs memory remains unclear. Here, a total of 42 C57BL6/J mice that exposed to no light at night, dim white light at night (dLAN-WL), or dim blue light at night (dLAN-BL) for 28 days. Behavioral data indicated that exposure to dLAN-BL resulted in severe recognition memory impairment, as evidenced by the reduced recognition index and discrimination index in the novel object recognition test. At the same time, we observed a decrease in plasma insulin levels. Consistent with these changes, we also observed that dLAN-BL reduced the number of neurons in the CA1, CA3 and DG regions of the hippocampus, up-regulated the mRNA expression levels of Bax, down-regulated the mRNA expression levels of Bcl-2, Bcl-xl and the protein expression level of pIRS1, pAKT, pGSK3ß, ß-catenin in the hippocampus. In vitro experiments, we found that insulin (10 nM) inhibited apoptosis and up-regulated the protein expression levels of pAKT, pGSK3ß, ß-catenin of HT22 cells induced by H2O2 (200 µM). However, these changes disappeared when the insulin receptors (IR) in HT22 cells were silenced. Taken together, our findings suggested that the impairment of memory in mice induced by dLAN-BL was mediated by insulin via the IR/IRS1/AKT/GSK3ß/ß-catenin pathway. DATA AVAILABILITY: All data generated or analyzed during this study are included in this published article.

An Autonomous City-Wide Light Pollution Measurement Network System Using LoRa Wireless Communication.

Light pollution is an ongoing problem for city populations. Large numbers of light sources at night negatively affect humans' day-night cycle. It is important to measure the amount of light pollution in order to effectively ascertain the amount of light pollution in the city area and effectively reduce it where possible and necessary. In order to perform this task, a prototype wireless sensor network for automated, long-term measurement of light pollution was developed for the Torun (Poland) city area. The sensors use LoRa wireless technology to collect sensor data from an urban area by way of networked gateways. The article investigates the sensor module architecture and design challenges as well as network architecture. Example results of light pollution measurements are presented, which were obtained from the prototype network.

Aerosol impact on light pollution in cities and their environment.

Measurements of artificial light at night represent an incredible challenge as the optical state of the atmosphere is highly unstable thus making both long-term trend analyses and inter-comparison of multiple observations difficult. Variations of atmospheric parameters, caused by either natural or anthropogenic processes, can massively influence the level of resulting night sky brightness caused by light pollution. Focusing on six parameters, either from aerosol optics or emission properties of light sources, this work literarily and numerically examines defined variations in aerosol optical depth, asymmetry parameter, single scattering albedo, ground surface reflectance, direct uplight ratio, and aerosol scale height. For each individual element the effect size and angular reliance is investigated, with results indicating that besides the aerosol scale height all play non-negligible roles in forming skyglow and environmental impact. Especially variations in aerosol optical depth and city emission function displayed severe discrepancies in consequential light pollution level. Hence, future improvement on atmospheric condition, i.e., air quality, focusing particularly on discussed elements indicates to positively influence the level of environmental impact caused by artificial light at night. We underline the need of inclusion of our outcomes to urban development and civil engineering processes in order to create or protect habitable areas for humans, wildlife and nature.