28Human activities have caused a near-ubiquitous and evolutionarily-unprecedented increase in 29 environmental sound levels and artificial night lighting. These stimuli reorganize communities 30 by interfering with species-specific perception of time cues, habitat features, and auditory and 31 visual signals. Rapid evolutionary changes could occur in response to light and noise, given their 32 magnitude, geographical extent, and degree to which they represent unprecedented 33 environmental conditions. We present a framework for investigating anthropogenic light and 34 noise as agents of selection, and as drivers of other evolutionary processes, to influence a range 35 of behavioural and physiological traits, such as phenological characters and sensory and 36 signalling systems. In this context, opportunities abound for understanding contemporary and 37 rapid evolution in response to human-caused environmental change. The overcast night sky radiance in urban areas has been found to be as much as four orders of 55 magnitude larger than in natural settings (Figure 1) [5]. Similarly, increased noise levels affect a 56 sizable proportion of the human population. In Europe for instance, 65% of the population is 57 exposed to ambient sound levels exceeding 55 dB(A) [6], roughly equivalent to constant rainfall. 58Of the land in the contiguous U.S., 88% is estimated to experience elevated sound levels from 59 anthropogenic noise (Figure 1) [7]. These effects are not limited to terrestrial environments; 60 ocean noise levels are estimated to have increased by 12 decibels (an ~16-fold increase in sound 61 intensity) in the past few decades from commercial shipping alone [8], while an estimated 22% 62 of the global coastline is exposed to artificial light [3] and many offshore coral reefs are 63 chronically exposed to artificial lighting from cities, fishing boats, and hydrocarbon extraction 64 [9]. 65The changes in light at night and noise levels are occurring on a global scale similar to 66 well-recognized ecological and evolutionary forces such as land cover and climate change. In 67 4 parallel with research involving climate change [10], much of our understanding of organismal 68 response to noise and light is restricted to short-term behavioural reactions. Organismal 69 responses might be associated with tolerance to these stimuli in terms of habitat use [11,12] Status of research on anthropogenic light and sound in ecology 98Night lighting and noise are highly correlated in many landscapes (e.g., [21]). It is critical to 99 understand whether the selective pressures these stimuli exert are additive, synergistic (Figure 2), 100 or if they mitigate one another. Few studies have examined the influence of each simultaneously 101 (e.g., [21]). In one study, flashing lights combined with boat motor noise suppressed antipredator 102 behaviour in hermit crabs (Coenobita clypeatus) more so than noise alone [22]. Future research 103 should quantify both light and sound simultaneously in the same population. Existing r...

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Phase and period responses of the circadian system of mice (Mus musculus) to light stimuli of different duration Comas, M.; Beersma, D. G. M.; Spoelstra, K.; Daan, S. Take-down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim.Downloaded from the University of Groningen/UMCG research database (Pure): http://www.rug.nl/research/portal. For technical reasons the number of authors shown on this cover page is limited to 10 maximum. In nature, virtually all circadian rhythms assume the 24.0-h period of the solar day-night cycle. This is due to the entrainment of the endogenous oscillators to the external light-dark cycle, the dominant zeitgeber for the majority of organisms. The process of entrainment is based on differential phase and period responses of the circadian systems to light depending on the phase at which the stimulus is applied. The (1, 3, 4, 6, 9, 12, and 18 h) given once per 11 days in otherwise constant darkness. Light-pulse duration affected both amplitude and shape of the phase response curve. Nine-hour light pulses yielded the maximal amplitude PRC. As in other systems, the circadian period slightly lengthened following delays and shortened following advances. The authors aimed to understand how different parts of the light signal contribute to the eventual phase shift. When PRCs were plotted using the onset, midpoint, and end of the pulse as a phase reference, they corresponded best with each other when using the mid-pulse. Using a simple phase-only model, the authors explored the possibility that light affects oscillator velocity strongly in the 1st hour and at reduced strength in later hours of the pulse due to photoreceptor adaptation. They fitted models based on the 1-h PRC to the data for all light pulses. The best overall correspondence between PRCs was obtained when the effect of light during all hours after the first was reduced by a factor of 0.22 relative to the 1st hour. For the predicted PRCs, the light action centered on average at 38% of the light pulse. This is close to the reference phase yielding best correspondence at 36% of the pulses. The result is thus compatible with an initial major contribution of the onset of the light pulse followed by a reduced effect of light responsible for the differences between PRCs for different duration pulses. The authors suggest that the mid-pulse is a better phase reference than lights-on to plot and compare PRCs of different light-pulse durations.

Artificial night-time illumination of natural habitats has increased dramatically over the past few decades. Generally, studies that assess the impact of artificial light on various species in the wild make use of existing illumination and are therefore correlative. Moreover, studies mostly focus on short-term consequences at the individual level, rather than long-term consequences at the population and community level—thereby ignoring possible unknown cascading effects in ecosystems. The recent change to LED lighting has opened up the exciting possibility to use light with a custom spectral composition, thereby potentially reducing the negative impact of artificial light. We describe here a large-scale, ecosystem-wide study where we experimentally illuminate forest-edge habitat with different spectral composition, replicated eight times. Monitoring of species is being performed according to rigid protocols, in part using a citizen-science-based approach, and automated where possible. Simultaneously, we specifically look at alterations in behaviour, such as changes in activity, and daily and seasonal timing. In our set-up, we have so far observed that experimental lights facilitate foraging activity of pipistrelle bats, suppress activity of wood mice and have effects on birds at the community level, which vary with spectral composition. Thus far, we have not observed effects on moth populations, but these and many other effects may surface only after a longer period of time.

Circadian rhythms with an endogenous period close to or equal to the natural light-dark cycle are considered evolutionarily adaptive ("circadian resonance hypothesis"). Despite remarkable insight into the molecular mechanisms driving circadian cycles, this hypothesis has not been tested under natural conditions for any eukaryotic organism. We tested this hypothesis in mice bearing a short-period mutation in the enzyme casein kinase 1e (tau mutation), which accelerates free-running circadian cycles. We compared daily activity (feeding) rhythms, survivorship, and reproduction in six replicate populations in outdoor experimental enclosures, established with wild-type, heterozygous, and homozygous mice in a Mendelian ratio. In the release cohort, survival was reduced in the homozygote mutant mice, revealing strong selection against shortperiod genotypes. Over the course of 14 mo, the relative frequency of the tau allele dropped from initial parity to 20%. Adult survival and recruitment of juveniles into the population contributed approximately equally to the selection for wild-type alleles. The expression of activity during daytime varied throughout the experiment and was significantly increased by the tau mutation. The strong selection against the short-period tau allele observed here contrasts with earlier studies showing absence of selection against a Period 2 (Per2) mutation, which disrupts internal clock function, but does not change period length. These findings are consistent with, and predicted by the theory that resonance of the circadian system plays an important role in individual fitness.circadian rhythms | survival | reproduction | resonance | tau mutation C ircadian clocks are a ubiquitous feature of life on earth, and serve to maintain synchrony of internal physiology with the external 24-h environment. Colin Pittendrigh, one of the founders of chronobiology, hypothesized that natural selection should favor circadian systems to operate in resonance with the external cycle (1, 2). A prediction from this hypothesis is that individuals exhibiting circadian rhythms with frequencies that are not in close resonance with the 24-h cycle should be selected against in nature. The hypothesis was initially supported by laboratory experiments in fly species that lived longer in a 24-h light-dark (LD) cycle than in non-24-h LD cycles (2-4). Stronger support emerged from dyadic competition experiments in batch cultures of cyanobacteria carrying single gene mutations affecting their circadian period (τ). Strains (either wild type or mutant) with a τ similar to the external LD cycle outcompeted strains with a τ different from the Zeitgeber (5, 6). Whether periods out of resonance with the external cycle entail a real fitness deficit in a natural setting has not been tested in any of these systems.The Ck1e tau (hereafter defined as the tau mutation) is a gain-offunction mutation (7) that accelerates the cellular dynamics of the circadian PERIOD protein (8, 9) and affects circadian behavior and physiology (10). It was f...

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.

The daily recurrence of activity and rest are so common as to seem trivial. However, they reflect a ubiquitous temporal programme called the circadian clock. In the absence of either anatomical clock structures or clock genes, the timing of sleep and wakefulness is disrupted. The complex nature of circadian behaviour is evident in the fact that phasing of the cycle during the day varies widely for individuals, resulting in extremes colloquially called 'larks' and 'owls'. These behavioural oscillations are mirrored in the levels of physiology and gene expression. Deciphering the underlying mechanisms will provide important insights into how the circadian clock affects health and disease. Keywords: circadian rhythm; entrainment; phase; oscillator; clock EMBO reports (2005) 6, 930-935.

We tested the hypotheses that the distance bats fly from tree lines depend on food abundance and protection from wind. We monitored the activity of pipistrelle bats (Pipistrellus pipistrellus) and measured insect abundance and wind speed and direction at fixed distances up to 50 m from tree lines. We compared bat behaviour in different situations: with and without wind and with low and high insect abundances in adjacent open areas. In all situations, pipistrelle bats' activity decreased with increasing distance from the tree line. Within nights, we found no effect of wind speed on bat activity (sound recorded per 5 min) on the leeward side of the tree lines. Between nights, however, bats concentrated their activities closer to the tree lines at high wind speeds or angles of incidence of wind from 45° to 90°. A significant relationship between bat and insect abundances was found only when the tree line was bordered by insect-rich grassland. Since wind and insect abundance only partly explained the distances bats flew from tree lines, two alternative explanations, namely predator avoidance and the use of tree lines as acoustic landmarks, are discussed. Pipistrelle bats using a double row of trees as a commuting route at dusk flew mainly between the tree lines, regardless of insect abundance or wind speed. It is argued that predator avoidance explains this behaviour, being a constraint on movements of bats at relatively high light levels. At high wind speeds and angles of incidence greater than 45°, the proportion of pipistrelle bats commuting on the leeward side of the tree lines increased.