Potential Biological and Ecological Effects of Flickering Artificial Light

Organisms have evolved under stable natural lighting regimes, employing cues from these to govern key ecological processes. However, the extent and density of artificial lighting within the environment has increased recently, causing widespread alteration of these regimes. Indeed, night-time electric lighting is known significantly to disrupt phenology, behaviour, and reproductive success, and thence community composition and ecosystem functioning. Until now, most attention has focussed on effects of the occurrence, timing, and spectral composition of artificial lighting. Little considered is that many types of lamp do not produce a constant stream of light but a series of pulses. This flickering light has been shown to have detrimental effects in humans and other species. Whether a species is likely to be affected will largely be determined by its visual temporal resolution, measured as the critical fusion frequency. That is the frequency at which a series of light pulses are perceived as a constant stream. Here we use the largest collation to date of critical fusion frequencies, across a broad range of taxa, to demonstrate that a significant proportion of species can detect such flicker in widely used lamps. Flickering artificial light thus has marked potential to produce ecological effects that have not previously been considered.     

Restless roosts: Light pollution affects behavior,sleep, and physiology in a free‐living songbird

The natural nighttime environment is increasingly polluted by artificial light. Several studies have linked artificial light at night to negative impacts on human health. In free‐living animals, light pollution is associated with changes in circadian, reproductive, and social behavior, but whether these animals also suffer from physiologic costs remains unknown. To fill this gap, we made use of a unique network of field sites which are either completely unlit (control), or are artificially illuminated with white, green, or red light. We monitored nighttime activity of adult great tits, Parus major, and related this activity to within‐individual changes in physiologic indices. Because altered nighttime activity as a result of light pollution may affect health and well‐being, we measured oxalic acid concentrations as a biomarker for sleep restriction, acute phase protein concentrations and malaria infection as indices of immune function, and telomere lengths as an overall measure of metabolic costs. Compared to other treatments, individuals roosting in the white light were much more active at night. In these individuals, oxalic acid decreased over the course of the study. We also found that individuals roosting in the white light treatment had a higher probability of malaria infection. Our results indicate that white light at night increases nighttime activity levels and sleep debt and affects disease dynamics in a free‐living songbird. Our study offers the first evidence of detrimental effects of light pollution on the health of free‐ranging wild animals.     

Multiple night‐time light‐emitting diode lighting strategies impact grassland invertebrate assemblages

White light‐emitting diodes (LEDs) are rapidly replacing conventional outdoor lighting technologies around the world. Despite rising concerns over their impact on the environment and human health, the flexibility of LEDs has been advocated as a means of mitigating the ecological impacts of globally widespread outdoor night‐time lighting through spectral manipulation, dimming and switching lights off during periods of low demand. We conducted a three‐year field experiment in which each of these lighting strategies was simulated in a previously artificial light naïve grassland ecosystem. White LEDs both increased the total abundance and changed the assemblage composition of adult spiders and beetles. Dimming LEDs by 50% or manipulating their spectra to reduce ecologically damaging wavelengths partially reduced the number of commoner species affected from seven to four. A combination of dimming by 50% and switching lights off between midnight and 04:00 am showed the most promise for reducing the ecological costs of LEDs, but the abundances of two otherwise common species were still affected. The environmental consequences of using alternative lighting technologies are increasingly well established. These results suggest that while management strategies using LEDs can be an effective means of reducing the number of taxa affected, averting the ecological impacts of night‐time lighting may ultimately require avoiding its use altogether.     

Pollination by nocturnal Lepidoptera,and the effects of light pollution: a review

1. Moths (Lepidoptera) are the major nocturnal pollinators of flowers. However, their importance and contribution to the provision of pollination ecosystem services may have been under‐appreciated. Evidence was identified that moths are important pollinators of a diverse range of plant species in diverse ecosystems across the world. 2. Moth populations are known to be undergoing significant declines in several European countries. Among the potential drivers of this decline is increasing light pollution. The known and possible effects of artificial night lighting upon moths were reviewed, and suggest how artificial night lighting might in turn affect the provision of pollination by moths. The need for studies of the effects of artificial night lighting upon whole communities of moths was highlighted. 3. An ecological network approach is one valuable method to consider the effects of artificial night lighting upon the provision of pollination by moths, as it provides useful insights into ecosystem functioning and stability, and may help elucidate the indirect effects of artificial light upon communities of moths and the plants they pollinate. 4. It was concluded that nocturnal pollination is an ecosystem process that may potentially be disrupted by increasing light pollution, although the nature of this disruption remains to be tested.     

Why artificial light at night should be a focus for global change research in the 21st century

The environmental impacts of artificial light at night have been a rapidly growing field of global change science in recent years. Yet, light pollution has not achieved parity with other global change phenomena in the level of concern and interest it receives from the scientific community, government and nongovernmental organizations. This is despite the globally widespread, expanding and changing nature of night‐time lighting and the immediacy, severity and phylogenetic breath of its impacts. In this opinion piece, we evidence 10 reasons why artificial light at night should be a focus for global change research in the 21st century. Our reasons extend beyond those concerned principally with the environment, to also include impacts on human health, culture and biodiversity conservation more generally. We conclude that the growing use of night‐time lighting will continue to raise numerous ecological, human health and cultural issues, but that opportunities exist to mitigate its impacts by combining novel technologies with sound scientific evidence. The potential gains from appropriate management extend far beyond those for the environment, indeed it may play a key role in transitioning towards a more sustainable society.     

Direct and indirect effects of light pollution on the performance of an herbivorous insect

Light pollution is a global disturbance with resounding impacts on a wide variety of organisms, but our understanding of these impacts is restricted to relatively few higher vertebrate species. We tested the direct effects of light pollution on herbivore performance as well as indirect effects mediated by host plant quality. We found that artificial light from streetlights alters plant toughness. Additionally, we found evidence of both direct and indirect effects of light pollution on the performance of an herbivorous insect, which indicates that streetlights can have cascading impacts on multiple trophic levels. Our novel findings suggest that light pollution can alter plant-insect interactions and thus may have important community-wide consequences.     

Light pollution alters the phenology of dawn and dusk singing in common European songbirds

Artificial night lighting is expanding globally, but its ecological consequences remain little understood. Animals often use changes in day length as a cue to time seasonal behaviour. Artificial night lighting may influence the perception of day length, and may thus affect both circadian and circannual rhythms. Over a 3.5 month period, from winter to breeding, we recorded daily singing activity of six common songbird species in 12 woodland sites, half of which were affected by street lighting. We previously reported on analyses suggesting that artificial night lighting affects the daily timing of singing in five species. The main aim of this study was to investigate whether the presence of artificial night lighting is also associated with the seasonal occurrence of dawn and dusk singing. We found that in four species dawn and dusk singing developed earlier in the year at sites exposed to light pollution. We also examined the effects of weather conditions and found that rain and low temperatures negatively affected the occurrence of dawn and dusk singing. Our results support the hypothesis that artificial night lighting alters natural seasonal rhythms, independently of other effects of urbanization. The fitness consequences of the observed changes in seasonal timing of behaviour remain unknown.     

Artificial night lighting disrupts sex pheromone in a noctuid moth

1. One major, yet poorly studied, change in the environment is the increase in nocturnal light pollution. Although this strongly alters the habitat of nocturnal species, the ecological consequences are poorly known. Moths are well known to be attracted to artificial light sources, but artificial light may affect them in other ways as well. 2. In this study, female Mamestra brassicae moths were subjected to various types of low‐intensity artificial night lighting with contrasting spectral compositions (green‐rich, red‐rich, warm white) or to a dark control treatment and the effects on their sex pheromone production and composition were tested. 3. Artificial night lighting reduced sex pheromone production and altered the chemical composition of the pheromone blend, irrespective of spectral composition. Specifically, amounts of the main pheromone component Z11‐16:Ac were reduced, while the deterring compounds Z9‐14:Ac, Z9‐16:Ac, and Z11‐16:OH were increased relative to Z11‐16:Ac when females were kept under artificial light. These changes may reduce the effectiveness of the sex pheromones, becoming less attractive for males. 4. These results show for the first time that artificial light at night affects processes that are involved in moth reproduction. The potential for mitigation through manipulation of the spectral composition of artificial light appears limited.     

Mate choice behavior of female Eastern Gray Treefrogs (Hyla versicolor) is robust to anthropogenic light pollution

Human activities are drastically changing the amount of artificial light entering natural habitats. Because light pollution alters the sensory environment, it may interfere with behaviors ranging from prey detection and vigilance to mate choice. Here, we test the hypothesis that anthropogenic light pollution affects the mate choice behavior of female Eastern Gray Treefrogs (Hyla versicolor). We tested this hypothesis under two experimental light treatments that simulate the light pollution created by streetlights (expansion of lit areas and increased light intensity), and the light pollution created by headlights of passing vehicles (rapid fluctuations between bright and dark conditions). The hypothesis predicts that females tested under conditions simulating light pollution will show behavioral changes geared toward mitigating detection by predators, such as relaxed preferences, decreased choosiness for the normally preferred call, and differences in approach behavior (either more directional, faster, or stealthier movements, or no approach at all). Contrary to our prediction, we found that light pollution did not affect mate choice behavior in Gray Treefrogs, and should therefore neither interfere with population persistence nor affect the sexual selection regimes on male call traits of this species. However, we caution that this result does not imply that anthropogenic light pollution is of no concern for amphibian conservation, because behavioral responses to variation in nocturnal light levels (both in the natural as well as anthropogenically enhanced range) seem to be highly species‐specific in anurans. We encourage additional studies to help gage the vulnerability of anurans to anthropogenic light pollution.     

Spectral properties of light affect plasma and pituitary gonadotropins in male rats

In order to find out whether different light spectra have any role in regulating the gonadotropin levels in male rats, we compared the 24-hour patterns of plasma and pituitary gonadotropins in rats kept for 7 days in natural or in cool white artificial lighting (exp. I). The intensity and periodicity of the two lighting conditions were adjusted as similar as possible. Further, we measured plasma and pituitary gonadotropins in the middle of the light period and in the middle of the dark period in rats kept for 7 days under artificial lightings of three different spectra (exp. II). In both experiments, in all lighting conditions we found higher plasma levels of LH and FSH during the dark than the light period. The differences were statistically significant only when the illumination contained more long and/or short wavelengths than the usual cool white laboratory lighting. The pituitary contents of gonadotropins were not found to vary with the periodicity of lighting. In the 24-hour patterns the overall plasma levels were higher and the pituitary contents of gonadotropins lower in natural lighting than in cool white lighting. It was concluded that the spectral properties of light influence the secretion of gonadotropins in male rats, but the mechanism involved remains to be clarified.     

Remote sensing to map influence of light pollution on Cory’s shearwater in São Miguel Island,Azores Archipelago

Global economic and population growth increase the extent and intensity of artificial night lighting. From an ecological perspective, this is light pollution, which causes changes in reproductive physiology, migration and foraging of many species and ultimately leads to loss of biodiversity. Some seabirds are intimately linked with the light features of their environments because they are nocturnally active. We report light-induced groundings of Cory’s shearwater (Calonectris diomedea) during a 2-year study (2008 and 2009) in São Miguel Island, in the Azores archipelago, and investigate the spatial correlation of locations of grounded birds with an annual composite of remotely sensed stable lights. Results indicate that 16.7% of fledglings are attracted to lights. The exposure of shearwater colonies in the study area to artificial night lighting is low overall. Four colonies account for 87% of the grounded birds. The distance each bird was found from the closest colony was best explained by the ratio of the satellite-measured light levels at the grounding spot to the light levels at the assigned colony of origin. These results demonstrate that satellite-observed nighttime lights are sufficient to assess risk to marine birds at the scale of oceanic islands and indicate their utility for monitoring the effectiveness of programs to manage lighting to reduce risk for these species and conducting global assessments of species vulnerability. To minimize the impact on Cory’s shearwater and other marine birds, we recommend measures such as reduction and control of lighting intensity near colony locations, while continuing and re-enforcing rescue campaigns.     

Light pollution as an ecological edge effect: Landscape ecological analysis of light pollution in protected areas in Korea

Protected areas have been increasingly encroached upon by light pollution. This paper examines whether patch geometry and land development intensities in protected areas affect the light pollution levels in 22 Korean national parks. Light pollution levels were assessed based on mean nighttime radiance of these national parks derived from nighttime satellite images. Total floor areas of buildings in the parks and in a 1 km buffer zone outside the park boundaries were calculated to assess land development intensities both inside and surrounding the parks. Two landscape metrics (patch area and fractal dimension) were measured to assess the patch geometry of the parks. Results show that the mean nighttime radiances in the Korean national parks are significantly affected by total floor areas in the 1 km buffer zone, but not by the total floor area within the park. This suggests that skyglow caused by artificial light in periphery is a major source of light pollution in the Korean national parks. In other words, light pollution can be seen as an ecological edge effect in protected areas where land development is stringently restricted. This conclusion is also supported by significant effects of two patch geometry variables, i.e., light pollution was more severe in a protected area with smaller size and a more complex shape. Based on these findings, it is suggested to make protected areas larger and more simple-shaped to protect biodiversity from light pollution. Managing lights in periphery of protected areas is also suggested.     

光和噪声污染胁迫下城市生态斑块鸟类风险评价

为应对城市生态斑块光污染、噪声污染范围快速扩张和强度增加对鸟类带来的生态风险,探讨了将微观鸟类风险阈值与宏观光污染、噪声污染分布数据结合进行生态风险评价的全链条研究方法,具体包括：探究微观层面光、声污染胁迫下鸟类生态风险阈值,分别获得光污染对黄雀和栗鹀寤寐节律及声污染对画眉鸟退避行为的生态风险阈值,并提出由实验室环境中获得的鸟类光污染强度风险阈值推算室外夜间混合光环境中鸟类光污染强度风险阈值的方法;同时获取宏观光、声环境分布数据,其中光环境分布数据基于Luojia 1-01和Jilin 1-7B夜光遥感影像获得,声环境分布数据通过软件模拟、实测校核、ArcGIS属性赋予等系列操作获得;结合微观鸟类风险阈值和宏观光、声环境分布数据开展生态风险评价,以典型城市生态斑块为实例分析光污染和噪声污染胁迫下鸟类生态风险分布特征;搭建生态风险评价平台并进行鸟类生态风险可视化展现。该生态风险评价方法可为城市宏观区域光、声污染生态风险的快速评价提供科学规范的研究和技术范例。     

Conserving energy at a cost to biodiversity? Impacts of LED lighting on bats

Artificial lighting is a key biodiversity threat and produces 1900 million tonnes of CO ₂ emissions globally, more than three times that produced by aviation. The need to meet climate change targets has led to a global increase in energy‐efficient light sources such as high‐brightness light‐emitting diodes (LEDs). Despite the energetic benefits of LEDs, their ecological impacts have not been tested. Using an experimental approach, we show that LED street lights caused a reduction in activity of slow‐flying bats ( Rhinolophus hipposideros and Myotis spp.). Both R. hipposideros and Myotis spp. activities were significantly reduced even during low light levels of 3.6 lux. There was no effect of LED lighting on the relatively fast‐flying Pipistrellus pipistrellus, Pipistrellus pygmaeus and Nyctalus/Eptesicus spp. We provide the first evidence of the effects of LED lights on bats. Despite having considerable energy‐saving benefits, LED lights can potentially fragment commuting routes for bats with associated negative conservation consequences. Our results add to the growing evidence of negative impacts of lighting on a wide range of taxa. We highlight the complexities involved in simultaneously meeting targets for reduction of greenhouse gas emissions and biodiversity loss. New lighting strategies should integrate climate change targets with the cultural, social and ecological impacts of emerging lighting technologies.     

Is part‐night lighting an effective measure to limit the impacts of artificial lighting on bats?

As light pollution is currently considered to be a major threat to biodiversity, different lighting management options are being explored to mitigate the impact of artificial lighting on wildlife. Although part‐night lighting schemes have been adopted by many local authorities across Europe to reduce the carbon footprint and save energy, their effects on biodiversity are unknown. Through a paired, in situ experiment, we compared the activity levels of 8 bat species under unlit, part‐night, and full‐night lighting treatments in a rural area located 60 km south of Paris, France. We selected 36 study locations composed of 1 lit site and a paired unlit control site; 24 of these sites were located in areas subject to part‐night lighting schemes, and 12 sites were in areas under standard, full‐night lighting. There was significantly more activity on part‐night lighting sites compared to full‐night lighting sites for the late‐emerging, light‐sensitive Plecotus spp., and a similar pattern was observable for Myotis spp., although not significant. In contrast, part‐night lighting did not influence the activity of early emerging bat species around streetlights, except for Pipistrellus pipistrellus for which there was significantly less activity on part‐night lighting sites than on full‐night lighting sites. Overall, no significant difference in activity between part‐ and full‐night lighting sites were observed in 5 of the 8 species studied, suggesting that current part‐night lighting schemes fail to encompass the range of activity of most bat species. We recommend that such schemes start earlier at night to effectively mitigate the adverse effects of artificial lighting on light‐sensitive species, particularly along ecological corridors that are especially important to the persistence of biodiversity in urban landscapes.     

Night-time lighting alters the composition of marine epifaunal communities

Marine benthic communities face multiple anthropogenic pressures that compromise the future of some of the most biodiverse and functionally important ecosystems in the world. Yet one of the pressures these ecosystems face, night-time lighting, remains unstudied. Light is an important cue in guiding the settlement of invertebrate larvae, and altering natural regimes of nocturnal illumination could modify patterns of recruitment among sessile epifauna. We present the first evidence of night-time lighting changing the composition of temperate epifaunal marine invertebrate communities. Illuminating settlement surfaces with white light-emitting diode lighting at night, to levels experienced by these communities locally, both inhibited and encouraged the colonization of 39% of the taxa analysed, including three sessile and two mobile species. Our results indicate that ecological light pollution from coastal development, shipping and offshore infrastructure could be changing the composition of marine epifaunal communities.     

Artificial light at night causes diapause inhibition and sex‐specific life history changes in a moth

Rapidly increasing levels of light pollution subject nocturnal organisms to major alterations of their habitat, the ecological consequences of which are largely unknown. Moths are well‐known to be attracted to light at night, but effects of light on other aspects of moth ecology, such as larval development and life‐history, remain unknown. Such effects may have important consequences for fitness and thus for moth population sizes. To study the effects of artificial night lighting on development and life‐history of moths, we experimentally subjected Mamestra brassicae (Noctuidae) caterpillars to low intensity green, white, red or no artificial light at night and determined their growth rate, maximum caterpillar mass, age at pupation, pupal mass and pupation duration. We found sex‐specific effects of artificial light on caterpillar life‐history, with male caterpillars subjected to green and white light reaching a lower maximum mass, pupating earlier and obtaining a lower pupal mass than male caterpillars under red light or in darkness. These effects can have major implications for fitness, but were absent in female caterpillars. Moreover, by the time that the first adult moth from the dark control treatment emerged from its pupa (after 110 days), about 85% of the moths that were under green light and 83% of the moths that were under white light had already emerged. These differences in pupation duration occurred in both sexes and were highly significant, and likely result from diapause inhibition by artificial night lighting. We conclude that low levels of nocturnal illumination can disrupt life‐histories in moths and inhibit the initiation of pupal diapause. This may result in reduced fitness and increased mortality. The application of red light, instead of white or green light, might be an appropriate measure to mitigate negative artificial light effects on moth life history.     

Cost reduction in the micropropagation of banana by using tubular skylights as source for natural lighting

Summary Daylight instead of artificial light was exploited for the in vitro culture of banana. Tubular skylights rediverted natural light into an interior enelosed room whereby each skylight, available for ca. US$600, could sufficiently illuminate an area of 3–5 m². The maintenance-free system allowed only a minimum of heat transfer and no cooling was necessary. The culture room required no electricity supply and under our conditions savings on costs for electricity of US$6 m⁻² wk⁻¹ were achieved, as compared to a standard growth room equipped with artificial lighting and controlled photoperiod and temperature regimes. Under natural light conditions, micropropagated plantlets were well developed at mean photosynthetic photon flux densities (PPFD) of 5–13 μmol m⁻² s⁻¹ and photoperiods of 9–14 h. Micropropagation rates were either the same or significantly higher than under artificial lighting. Single shoots on rooting medium showed some symptoms of etiolation, yet acclimatization rates averaged 95%. A step-like culture rack. rather than a vertical one, permitted uniform plant growth on all levels. This paper describes an established micropropagation system of low cost and simplicity.     

Impacts of “supermoon” events on the physiology of a wild bird

The position of the Moon in relation to the Earth and the Sun gives rise to several predictable cycles, and natural changes in nighttime light intensity are known to cause alterations to physiological processes and behaviors in many animals. The limited research undertaken to date on the physiological responses of animals to the lunar illumination has exclusively focused on the synodic lunar cycle (full moon to full moon, or moon phase) but the moon's orbit—its distance from the Earth—may also be relevant. Every month, the moon moves from apogee, its most distant point from Earth—and then to perigee, its closest point to Earth. Here, we studied wild barnacle geese (Branta leucopsis) to investigate the influence of multiple interacting lunar cycles on the physiology of diurnally active animals. Our study, which uses biologging technology to continually monitor body temperature and heart rate for an entire annual cycle, asks whether there is evidence for a physiological response to natural cycles in lunar brightness in wild birds, particularly “supermoon” phenomena, where perigee coincides with a full moon. There was a three‐way interaction between lunar phase, lunar distance, and cloud cover as predictors of nighttime mean body temperature, such that body temperature was highest on clear nights when the full moon coincided with perigee moon. Our study is the first to report the physiological responses of wild birds to “supermoon” events; the wild geese responded to the combination of two independent lunar cycles, by significantly increasing their body temperature at night. That wild birds respond to natural fluctuations in nighttime ambient light levels support the documented responses of many species to anthropogenic sources of artificial light, that birds seem unable to override. As most biological systems are arguably organized foremost by light, this suggests that any interactions between lunar cycles and local weather conditions could have significant impacts on the energy budgets of birds.     

Dim Light at Night Increases Immune Function in Nile Grass Rats,a Diurnal Rodent

With the widespread adoption of electrical lighting during the 20th century, human and nonhuman animals became exposed to high levels of light at night for the first time in evolutionary history. This divergence from the natural environment may have significant implications for certain ecological niches because of the important influence light exerts on the circadian system. For example, circadian disruption and nighttime light exposure are linked to changes in immune function. The majority of studies investigating the effects of light exposure and circadian disruption on the immune system use nocturnal rodents. In diurnal species, many hormones and immune parameters vary with secretion patterns 180° out of phase to those of nocturnal rodents. Thus, the authors investigated the effects of nighttime light exposure on immunocompetence in diurnal Nile grass rats (Arvicanthis niloticus). Rats were housed in either standard 14-h light (L):10-h dark (D) cycles with L ～150 lux and D 0 lux or dim light at night (dLAN) cycles of LD 14:10 with L ～150 lux and D 5 lux for 3 wks, then tested for plasma bactericidal capacity, as well as humoral and cell-mediated immune responses. Rats exposed to dLAN showed increased delayed-type hypersensitivity pinna swelling, which is consistent with enhanced cell-mediated immune function. dLAN rats similarly showed increased antibody production following inoculation with keyhole lymphocyte hemocyanin (KLH) and increased bactericidal capacity. Daytime corticosterone concentrations were elevated in grass rats exposed to nighttime dim light, which may have influenced immunological measures. Overall, these results indicate nighttime light affects immune parameters in a diurnal rodent. (Author correspondence: fonken.1@osu.edu)