Anthropogenic Light and Noise as Sensory Pollutants and New Selection Pressures

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There is no alternative way to receive information about our environment besides our senses, and only from our senses are we able to act in our environment. This is a consistent law of life on Earth, affecting the lives of each and every organism. In the time since humans have developed much of the world into metropolitan spaces, anthropogenic (human-made) sound and artificial light at night have risen to record levels [1]. Animals sharing these areas have been forced to adapt to these rapid changes or move out of the way completely. That isn’t just on land; aquatic habitats have also been stricken with these obstacles to survival as 22% of global coastline is exposed to artificial light and noise [1]. The influx of artificial light and noise qualifies as sensory pollution, resulting in conditions that make it more difficult for many species to properly receive information about the environment. Research on the neural basis of behavior in the field of neuroethology informs us on how much survival depends on visual and auditory sensory modalities, importantly allowing species to find food and mates or avoid predators. Superfluous light, or photopollution, and sound stimuli put selection pressures on behavioral traits, sensory systems, and physiology. Downstream outcomes of these pressures are extinction, changes to community structures, and deteriorated ecosystem cycles. It may be obvious the effects of these sensory pollutants has disrupted the natural flow of life and had a tremendous effect on biodiversity as a result of human presence. The fact that by 2008 only 10% of land on Earth was considered “remote” from urbanized areas [2] emphasizes the weight of the issues caused by human encroachment. These swift, inorganic alterations to the environment hasten natural selection to an unprecedented rate by adding new pressures. The speed of these changes make it unlikely that animals have the genetic variance to evolve in response to those conditions [1].

Sensory Pollutants: Light

Since the invention of electrical light over a century ago, urban development has vastly extended the presence of artificial light at night. While humans see this as a luxury of the 21st century or even take it for granted, the other species we share space with are dependent on natural light cycles for many behaviors vital to survival like knowing when to forage the most safely and effectively and when it’s safe to mate among other things. It’s estimated that more than 80% of the US population and the majority of the EU maintain nighttime brightness levels that surpass that of a full moon night [3]. In areas like those, light pollution creates a ‘sky glow’ by scattering artificial light into the atmosphere which ultimately disrupts natural light patterns. Normal light:dark cycles function to maintain circadian rhythms, mate choices, foraging habits, migration, and overall health outcomes.

Circadian Rhythms

The rise and fall of the sun each day is an environmental rhythm that organisms have evolved to be responsive to for their benefit. Circadian comes from a latin root to mean “around the day”. Light is an important environmental cue that organizes physiological rhythms, such as our biological clocks that regulate when we and other animals sleep, wake, and eat [4]. Constant presence of light through the day and night can alter production of hormones like melatonin [5], which can result in dysregulation of immune function and other hormonal pathways [6]. In connection with immune function, a constant light cycle can lead to oxidative stress [7] but production of melatonin can combat this with its role in generating powerful antioxidative effects [8]. The catch is that melatonin is produced in the absence of light, so photopollution impedes on this natural process.

Migration and Orientation

Migration is a vital act for survival and reproduction in many species. These events are usually seasonal movements of populations for resources or breeding purposes. It’s known that migratory patterns are driven by sky illumination across taxa [9- 13]. For this reason, migration can be considered a “circannual” rhythm, similar to circadian rhythms, because seasonal positions of celestial bodies can be used as cues to migrate but also to orient migratory species to their destinations [14-16, 10]. Effects normally modulated by natural lunar light levels can be functionally mimicked by even low levels of photopollution [17]. For salmonid fishes, the darkness of a new moon creates a hormone surge that causes them to migrate out of rivers and towards the ocean [18]. Avian migration is known to depend on melatonin for when they migrate and how they orient themselves [16,19]. Likewise, interference by unnatural light throws off circadian rhythms of monarch butterflies, which then obstructs their ability to orient on their migration [10]. Accordingly, migratory behavior is reliant on environmental rhythms or physiological cues that are modulated by light.

Photopollution can also attract nocturnally migrating birds and then disorient them along their journey, resulting in them being trapped in artificially lit areas [15]. This risk is exacerbated by the fact that they usually fly lower late at night and during bad weather, bringing them in closer proximity to the lights [17]. Once in artificially lit urban areas, birds fatally collide into buildings when they try to leave. Lighthouses, broadcast towers, boats, and oil platforms all contribute to mortality and disruption of migration routes due to the photopollution emitted [15, 20-22]. Not only birds, but other crucial nodes on ecosystems, such as dung beetles, are experiencing reduced fitness to photopollution. While dung beetles are vital for nutrient cycling of their ecosystems [23], they depend on the light of the Milky Way as an orientation marker and thus have lost this crucial cue [14]. Relatedly, dung beetle diversity has dropped in areas with increasing urbanization [24], but the direct cause of this change is still uncertain. This impedance on natural light levels that guide migration patterns is broader reaching than one would think, as sky glow brightens the sky as far as kilometers away from the light source [25]. That being the case, timing and efficacy of migration and orientation is indispensable for survival and reproduction for many species across taxa.

Foraging and Predation

Species’ daily rhythms have evolved to benefit functions of their daily lives. For instance, timeliness of decision to forage or mate need to account for predation risk at certain times of day. There are two primary hypotheses that address how foraging trends are impacted by light cycles. The Predation Risk Hypothesis [26] posits that the risk of being seen by a predator during certain lighting conditions affects how animals base their decisions on when to forage or mate. Not mutually exclusive is the Foraging Efficiency Hypothesis [27] that predicts that prey availability will be affected by levels of light and in turn impacts foraging payoffs. For instance, short eared owls have best foraging efficiency under the high illumination of full moon, while simultaneously their prey (deer mice) will reduce their foraging behavior in order to avoid being spotted by the owls during periods of high illumination [28]. It is for that very reason that desert and temperate rodents [28,29], fruit bats [30], sea birds [26], and many more also decrease foraging behavior under high lunar illumination. Therefore, since predator-prey interactions are dependent on light, they are altered as a result of photopollution. Any changes to the predator-prey relationships will have large-scale downstream effects on the ecological balance, as each species is a node in a densely connected web of life.

But could anthropogenic light ever help animals search for prey? It is also sometimes used to the benefit of foraging in certain predator-prey dynamics, usually in favor of diurnal, or daytime active, animals. Many animals that are usually active in the day, like some species of birds and reptiles for example, are able to exploit artificial light by continuing to effectively forage at nighttime because their visual systems are equipped for lit conditions [31,32]. At first this seems beneficial for diurnal animals, but the resulting change in ecological structure may not be favorable to them in the end. One study found that insects were most attracted to bluish-white lights and congregated around these types of lights in high numbers at night, so in turn, bats switched foraging habits to center more around these lights in order to earn the best possible meal [10]. The insects were attracted to yellowish lights as well, but in lower numbers and so bats less frequently hunted near those lights. This is evidence that bright street lamps, especially of high wavelength in the spectrum, increases the predation risk of many insects and consequently alters the survival of these species. If the insects species do not survive and propagate, what will the bats eat? It’s a cyclic system that needs to stay in balance for the sake of preserving biodiversity, and this is just one example of how that can get out of check.

Reproductive Strategies

Photopollution can also distort important light and timing cues involved in reproduction, such as mate choice, territoriality, nesting locations, and incubation. Female frogs are less choosy about their mates under higher lighting levels, as they need to get it done quickly so to not risk getting eaten [33]. For some birds like blacktailed godwits, they prefer nesting sites that are at least 300 meters away from artificial road lighting [17]. Nesting near photopollution comes at a cost for male mockingbirds, who will usually only sing after mating under full moon illumination unless they are situated by artificial light, in which case males will continuously sing when they normally wouldn’t be [34]. It’s also known to impede on incubation in other birds [35] and cause advanced spawning in fish [36]. These alterations to reproductive strategies certainly impacts brood success in unpredictable ways as the artificial lighting adds an extra selection pressure.

Overall Impact on Biodiversity

The unnatural presence of artificial light at night is a new and expanding selection pressure that has appeared within the past century, and its exigence is growing. Artificial skyglow suppresses lunar light cycles and obliterates the possibility of dark nights that are crucial for signalling animal behaviors [4,37]. Light emission is growing 3-6% every year on a global scale [38]. Since skyglows permeate kilometers out of their urban originating areas, these rearrangements of behavioral responses to light cycles could seriously change the distribution and action patterns of many species. Community interactions become imbalanced with changes to how light cycles affect connections among species, such as in competition and predation [17]. An example of this is daytime active animals that now continue their activity into the night [32]. A “perpetual full moon” due to photopollution is favorable for diurnal species, but not others. Nocturnal animals depend on dark conditions to forage and if this is compromised, they have increased predation risk and could go too hungry to survive [17]. Consequently, community structures will be reformed with the elimination of species that are a core nodes in the ecosystem, yet these effects are still being largely ignored.

Sensory Pollutants: Sound

Just as with light, sound is pertinent to many aspects of survival. Exposure to chronic noise as a result of growth of road networks, resource extraction, and other modes of human encroachment reduce animals’ ability to perceive acoustic signals in their environment. As they rely on the acoustic environment to forage, escape predators, and successfully reproduce, we can expect significant alterations to community structure and decreased fitness among species that most heavily rely on sound to survive. This does not only apply to land animals, but is present in aquatic environments as well. Transportation networks are one of the largest contributors to anthropogenic noise as 83% of the land area of the continental US was within 1000 meters of a road by 2003 [39], bringing average noise levels to an unnatural extent. The consequence of this is inhibition of sound perception by raising the threshold needed to hear important cues [40]. This effect, known as ‘masking’ in the discipline of bioacoustics, is expected to be growing with human urban development. With this in mind, anthropogenic noise qualifies as a sensory pollutant just as much as artificial light at night. Loud noises could cause negative physiological outcomes such as hearing loss [41], heightened stress [42], and hypertension [43]. Noisy human activities reshape animal ecology [44,45], such as with how species richness declines near roads and negatively correlated with traffic density [46,47]. As such, anthropogenic noise is a growing concern for animal conservation.

Communication

Impedance to the ability to discriminate one’s acoustic environment is a major disruptor of transference of signals between animals in a territory. The impact of anthropogenic noise on communication reaches broadly across taxa. Birds, primates, rodents, and cetaceans have adapted around the masking effect by changing their vocalizations  [48-51]. Calls can serve as alarms for animals to warn each other of nearby predators or even promote social cohesion [52-54]. Other types of signals baby birds’ food begging calls to their mother [55], primate contact calls [56], bat echolocation [57], as well as communication for reproduction in birds, cetaceans, and frogs [48-51]. Masking can lower the distance at which these calls are heard and thus weaken social networks and their benefits. While some frogs are able to adapt their calls to the frequency or duration needed to be heard in the presence of anthropogenic noise [58], other closely related frogs are not [59].

Communication is a significant factor in reproduction in many social animals. However, acoustic masking can get in the way of communication networks that enable songbirds to assess individuals for mate choice [60], making the future reproductive repercussions unpredictable as anthropogenic noise disrupts these networks [61]. Traffic noise can make it more difficult for female tungara frogs to locate males [62]. Within 100 meters of roads, songbirds are more likely to abandon nests or their young [63]. Ovenbird populations habitating near gasoline compressor stations had more naive first-time breeders and also experience reduced pairing success [64]. With background sound being an important habitat characteristic, it becomes clear that the presence of loud noise degradates territory quality and constraints animals’ ability to perceive important factors in their environment.

Foraging and Predation

Cues are not only important for communication, but detecting prey or predators as well. Sometimes sound is the only cue for food, such as in robins when they need to localize buried worms [65]. Rustling sounds are used by march hawks and lemus for finding prey [66,67]. The same goes for aquatic mammals, of which multiple types of dolphins [68-70] and sperm whale [71] were able to adapt their sound structures to compensate for anthropogenic noise, but others like humpback, beluga, and killer whales were not [72-74]. Those that couldn’t adapt had reduced foraging efficiency. On the other side of the predator-prey dynamic, individuals need to stay alert for specific sounds in their acoustic environment to avoid getting predated on. Tungara frogs specifically avoid the sound of their bat predator’s wingbeats [75] Some bird nestlings know to be quiet when they hear the footsteps of their main predator [76]. The excessive amounts of noise are usually perceived as predation risk, which provokes anti-predator tactics and discourages other behaviors importantly relevant to fitness. These effects can certainly predict population decline [44]. Soundscapes are highly pertinent and salient sources of information needed to survive. Anthropogenic disturbances to this keystone can therefore have severe consequences.

Physical and Behavioral Evolutionary Responses to Sensory Pollutants

The significance of natural light cues for fitness is more densely consequential than we can currently predict. Anthropogenic light and sound are not only sensory pollutants, but they’ve added unnatural selection pressures and have quickly become agents of selection. Just like other competitive pressures in the environment, they influence the success of certain behavioral and physiological traits, such as communication and sensory systems [1]. The rapid rise of these sensory pollutants is not giving organisms the time to even have the genetic variance that is can be an important aspect in order to properly adapt to new environmental conditions [1]. However, there are other ways animals are facing these new challenges.

In the immediate toolboxes of many species is flexibility in their behavior as they try to salvage their chances at survival and reproduction by altering their responses to light or sound cycle. This could be the decision to mate at an earlier time than usual to offset the risk of getting eaten [33], or songbirds continuing to sing past the full moon because artificial light increases the need to defend the nest [77]. These decisions still come at a cost, however, because it wastes energetic resources or can offset physiological rhythms.

There is also flexibility in the way individuals develop alongside the demands of their environment. For example, alteration of acoustic environment in the oceans can alter the developmental trajectories of marine vertebrates [78], insects [79], and rodents [80]. Likewise, some birds have been observed to incubate their nests longer under extended photoperiods, which can lead to a smaller hatchling size and directly affecting fitness to survive [81].

Large-scale changes across ecosystems or at the species-wide level relies on the smaller evolutionary changes relating to genes that best cope with the changes. Adaptive responses rely on the heritability of traits that can transfer fitness to deal with the changing environment through generations. Growth, reproduction [82,83] and survivorship [84] are all contingent on sensory systems like vision and hearing [1]. Preferential selection for traits of the visual system based on its acclimation to the amount of light in their habitats have been observed in not only mammals [85], but fish as well [86]. The kinds of sensory stimuli necessitated by the environment can bias the ways animals choose habitats or how they migrate, ultimately changing the structure of the gene pool in certain regions and populations. For instance, both unnatural light and sound modify patterns of settlement, dispersal, and migration patterns [4,17,87]. Some birds settled in noisy areas [88] and mammals have been observed to avoid highly lit areas [89] which have been causally linked to reduced population sizes and threatening the chance that populations have the genetic variance to adapt [1]. Each individual modification of behaviors responding to unnatural stimuli can be expected to have flow-on effects for the entire ecosystem [1].

Anthropogenic light and sound are not only sensory pollutants, but they’ve added unnatural selection pressures and have quickly become agents of selection.

Solutions and Future Directions

These sensory pollutants and selection pressures new to the environmental scene invoke great concern for biodiversity and conservation efforts. Hindered ability to adapt to a rapidly changing environment can alter ability to successfully mate or reproduce [90], elevate stress [91], and dampen the accuracy of anti-predator behavior [92]. Changes to these vital aspects of species’ survival create a domino effect on the balance of the entire ecosystem by modifying predator-prey relationships [88], nutrient cycling [24], or other essential aspects of ecological functioning.

Nevertheless, there is hope for ameliorating these effects. Longcore et al. [87] has suggested tuning the color spectrum of street lamps from blue-dominant, which attract insects and disturbs circadian rhythms, to less disruptive colors like red or orange. Hunter [93] advocates for careful urban planning that includes business parks or rooftop gardens that provide some plant and animal diversity to sustain populations in cities to make them into more flexible ecosystems. Legislators can also write and advocate for policy to regulate levels of human-caused artificial light and noise that could effectively diminish the impact of anthropogenic disturbances.

Needless to say, the myriad of effects of anthropogenic noise on the lives of animals and their instrumental roles in balancing their ecosystems is an exigent issue. The welfare of the animals we share space with is in our hands as we are the perpetrators of the instability of their environment. The time to act is now, before our actions quickly erase millions of years of evolutionary accomplishment.

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