Artificial light and biting flies: the parallel development of attractive light traps and unattractive domestic lights

Abstract

Light trapping is an important tool for monitoring insect populations. This is especially true for biting Diptera, where light traps play a crucial role in disease surveillance by tracking the presence and abundance of vector species. Physiological and behavioural data have been instrumental in identifying factors that influence dipteran phototaxis and have spurred the development of more effective light traps. However, the development of less attractive domestic lights has received comparatively little interest but could be important for reducing interactions between humans and vector insects, with consequences for reducing disease transmission. Here, we discuss how dipteran eyes respond to light and the factors influencing positive phototaxis, and conclude by identifying key areas for further research. In addition, we include a synthesis of attractive and unattractive wavelengths for a number of vector species. A more comprehensive understanding of how Diptera perceive and respond to light would allow for more efficient vector sampling as well as potentially limiting the risk posed by domestic lighting.

Graphical Abstract

Full text

Wilsonetal. Parasites Vectors (2021) 14:28
https://doi.org/10.1186/s13071-020-04530-3
REVIEW
Articial light andbiting ies: theparallel
development ofattractive light traps
andunattractive domestic lights
Roksana Wilson*, Andrew Wakefield, Nicholas Roberts and Gareth Jones
Abstract
Light trapping is an important tool for monitoring insect populations. This is especially true for biting Diptera, where
light traps play a crucial role in disease surveillance by tracking the presence and abundance of vector species. Physi-
ological and behavioural data have been instrumental in identifying factors that influence dipteran phototaxis and
have spurred the development of more effective light traps. However, the development of less attractive domestic
lights has received comparatively little interest but could be important for reducing interactions between humans
and vector insects, with consequences for reducing disease transmission. Here, we discuss how dipteran eyes respond
to light and the factors influencing positive phototaxis, and conclude by identifying key areas for further research. In
addition, we include a synthesis of attractive and unattractive wavelengths for a number of vector species. A more
comprehensive understanding of how Diptera perceive and respond to light would allow for more efficient vector
sampling as well as potentially limiting the risk posed by domestic lighting.
Keywords: Diptera, Light attraction, Phototaxis, Spectral wavelength preferences, Vector
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Background
Haematophagy (blood-feeding) has evolved indepen-
dently multiple times amongst the Diptera [1]. Haema-
tophagous flies are collectively known as ‘the biting flies’,
and their lifestyle facilitates the transmission of blood-
borne pathogens from one host animal to another. To
humans, the most dangerous and prolific of this group
are the mosquitoes (Culicidae), which cause hundreds of
thousands of deaths annually through the transmission of
pathogens such as malaria, dengue and yellow fever [2].
Other vectors include sand flies (Phlebotominae), black
flies (Simuliidae) and tsetse flies (Glossinidae), which
transmit pathogens that cause leishmaniasis, river-blind-
ness and African sleeping sickness, respectively—causing
disfigurement, disability and chronic suffering [2]. Bit-
ing midges (Ceratopogonidae) play a limited role in the
transmission of pathogens that cause disease in humans
but cause considerable economic impact by spreading
bluetongue virus and pathogens causing African horse
sickness amongst livestock [3]. ese diseases dispro-
portionately affect the poorest populations, with deaths
being highest in African countries [4].
e abundance and distribution of biting fly popula-
tions should be closely monitored so that the risk of dis-
ease outbreak can be determined and the effectiveness of
vector-control strategies evaluated. Light traps have been
criticised for their bias towards certain taxa and flies of
a certain parity status (particularly human-feeding, host-
seeking females), and catches can be unrepresentative of
the local population [5–8]. However, due to their wide-
spread availability, ease of use, lack of risk to collectors
from infectious flies and minimal influence from human
error, they are now routinely used in the capture of mos-
quitoes [9], midges [10] and sand flies [11]. Attention has
since shifted to the development of more attractive light
traps [12–16]. Highly attractive lights are more likely
Open Access
Parasites & Vectors
*Correspondence: roksana.wilson@bristol.ac.uk
School of Biological Sciences, University of Bristol, Life Sciences Building,
24 Tyndall Avenue, Bristol BS8 1TQ, UK
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Wilsonetal. Parasites Vectors (2021) 14:28
to detect rare vector species, or increase the capture of
sparsely distributed individuals, which allows for control
measures to be implemented when the risk of disease
outbreak is still low [17–19]. Larger catches also increase
the chances of finding infected flies, allowing for virus
isolation [18].
Despite the interest in light attraction, the role arti-
ficial lights play in facilitating disease transmission by
attracting vectors remains understudied [20]. e only
definitive example of this phenomenon so far is Chagas
disease, spread by triatomine bugs (Order: Hemiptera).
Proximity to street lights is linked to house infestation
by triatomines [21, 22], and a 2005 disease outbreak was
traced to sugarcane juice contaminated with triatomines
attracted to the lamp above the juice kiosk [20]. For dis-
eases transmitted by dipterans, correlations between
electrification and malaria have been reported in the
Solomon Islands [23], Burkina Faso [24], Uganda [25]
and Malawi [26]. However, determining whether these
outbreaks are caused by artificial lights attracting mos-
quitoes to human settlements requires further study.
Even so, as biting flies are attracted to light, the develop-
ment of less attractive domestic lights is of considerable
importance.
e aim of this review is to outline the ways artificial
lights can be made more or less attractive to mosqui-
toes, midges and sand flies and to identify areas where
further research is needed. By modifying light traps and
domestic lights, trapping efficiency for vectors could be
improved and the public health risk posed by electrifica-
tion could be reduced.
Physiology ofthedipteran eye
Neurophysiological studies on the visual systems of flies
provide a better understanding of light attraction as they
reveal sensitivity to different wavelengths of light. How-
ever, studies on biting flies are rare [27].
Ommatidia, the units making up the compound eye,
contain eight photoreceptor cells known as retinula
cells (labelled R1–R8). e spectral sensitivity of these
cells, i.e. which wavelength bands the receptor absorbs,
depends mainly on the visual pigment rhodopsin (Rh)
within each photoreceptor [28]. In Drosophila, the R1–
R6 cells express Rh1, which responds to a broad spectrum
of light, and thus these cells are believed to be achro-
matic. e R7 cell expresses Rh3 or Rh4 pigments (both
ultraviolet [UV] sensitive) and the R8 cell expresses Rh5
(blue sensitive) or Rh6 (green sensitive) pigments, and
these cells are assumed to be chromatic [29]. is UV/
blue/green sensitivity is highly conserved in insects [30].
For biting flies, spectral sensitivity data exist for Aedes
aegypti [31], Culex pipiens [32], Lutzomyia longipalpis
[33], Glossina morsitans [34], Tabanus nigrovittatus [35],
simuliid blackflies (species not provided) [36], Stomoxys
calcitrans [37] and Haematobia irritans [37]. ese taxa
all show dual peaks in sensitivity, with one peak in the
UV and another in the blue/green, and minimal sensitiv-
ity to longer wavelengths (Fig.1).
In insects, attraction to specific wavelength bands is
controlled by photoreceptors and post-receptor mecha-
nisms. Wild-type Drosophila prefer UV light over blue
and green wavelengths. However, blocking the activ-
ity of Dm8 neurons causes the flies to prefer green light
(525 nm) over UV (370 nm) [29]. Dm8 neurons are
wide-field amacrine cells located in the medulla. ey
are the post-synaptic targets of the UV-sensitive R7 cells
and provide lateral connections to neurons that pro-
ject to higher visual centres [29]. Similarly, silencing the
R1–R6 cells or the R7 and R8 cells causes flies to prefer
blue (430nm) over UV (350 nm) [38]. Wild-type Dros-
ophila also prefer blue wavelengths over green (565nm),
but inactivating the blue-sensitive Rh5, or removing the
UV-sensitive R7 cells, causes flies to prefer green [38].
e attractiveness of specific wavelength bands can also
vary throughout the day and appears to be circadian
regulated [39, 40]. Wild-type Drosophila show a peak of
UV (365nm) and blue (460nm) light avoidance behav-
iour during midday. However, mutant flies lacking in
cryptochrome (cry), the primary circadian light sensor
in Drosophila, exhibit a strong attraction to UV and blue
light at all times of the day [39]. ese null cry flies also
show an increased attraction to orange (595 nm) light
compared to wild-type control flies [39]. An in-depth
outline of the mechanisms underpinning phototaxis is
beyond the scope of this review, but species-specific dif-
ferences in wavelength preferences are likely a result of
subtle differences in neurophysiology.
Attraction to certain wavelengths is not necessarily
a sign of colour vision. True colour vision is the ability
to distinguish “light of different spectral compositions
(hues) independently of their intensities” [29]. It requires
at least two photoreceptors with different sensitivities,
and the neural framework to compare the outputs of
these receptors [41]. Flies possess at least two photore-
ceptor types and so fulfil the first precondition of colour
vision. However, it is disputed whether insects adapted
to low light conditions, which are those most attracted
by light traps, would possess colour vision [42]. ere
are subtle differences between the eyes of nocturnal
and diurnal flies, such as whether the eye is configured
to increase image resolution at the expense of light sen-
sitivity (apposition eye), increase light sensitivity at the
expense of resolution (optical superposition)—or a com-
bination of the two (neural superposition) [43]. ese dif-
ferences may affect how the fly perceives colour.
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Wilsonetal. Parasites Vectors (2021) 14:28
Factors thatinuence light attraction
Lighting technology
ere are three main types of lighting technology avail-
able to consumers for home use: incandescent; fluo-
rescent; and light-emitting diode (LED) (Fig. 2). e
earliest light bulbs were incandescent, and their glow
is the result of a wire filament being heated, although
most of the electrical current passing through the bulb
is emitted as infrared (IR) radiation (heat). Numerous
countries have issued bans on these bulbs due to this
extreme energy inefficiency [44]. Compact fluorescent
lamps (CFLs) became an ‘energy-saving’ alternative
to the incandescent bulb. ese lamps use mercury to
produce UV light, which is then converted into visible
light when it strikes the fluorescent phosphor coating
on the inside of the bulb. e amount of UV light con-
verted depends on the design or quality of the lamp,
and as mercury is a bioaccumulating pollutant, CFLs
are classified as hazardous waste during disposal. e
newest technology are the LEDs: semiconductors that
emit light when a current is passed through them. LEDs
have many advantages over other light types, including
improved energy efficiency, low power consumption,
longer lifetime, high durability, cheaper cost and the
ability to produce monochromatic light in a variety of
wavelengths [16].
Fig. 1 Electroretinograms (ERGs) showing the spectral sensitivities of a female Aedes aegypti [31], b female Culex pipiens [32], c female Lutzomyia
longipalpis [33], d Glossina morsitans [34], e male Simuliid blackflies [36], f young, female Tabanus nigrovittatus [35], g Stomoxys calcitrans [37] and
h female Haematobia irritans [37]. Figure is adapted from original publications [31–37]. Studies differ in their methods and specimens (age, sex,
chromatic adaptation, etc). Ultraviolet and blue/green wavelengths are highlighted in grey
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Wilsonetal. Parasites Vectors (2021) 14:28
e global market share of LED lamps was approxi-
mately 36% in 2015 and is predicted to reach 67–80% in
2022. e highest growth rates are expected in Africa,
Asia-Pacific, India and Latin America [45]. ese are
regions with high death rates from vector-borne dis-
ease [4], making it important to determine how these
lights perform as vector attractants. A small difference
in attractiveness between two lights can become a large
difference in insect numbers when many lights are used
over a wide area. Conversely, LEDs could represent a
cheap, easy and highly effective surveillance tool.
Light traps using incandescent bulbs generally catch
fewer flies than light traps emitting UV (<400nm) light
[46–52]. However, LEDs can be both less attractive [9,
15, 17, 49, 53–57] and more attractive [18, 19, 58–63]
than incandescent or fluorescent lights. In studies where
LEDs have been found to be less attractive, the LED is
usually white. Typically, white LEDs operate by convert-
ing almost all of the electrical energy they receive into
light in the visible spectrum. is means they lack the
IR (> 700nm) and UV peaks found in incandescent and
fluorescent lights, respectively (Fig.2). UV light has been
characterised as attractive to insects, and high amounts
of IR radiation could act as a thermal attractant along-
side the light [53, 56]. In contrast, LEDs emitting narrow
bands of short wavelength light generally attract more
insects than the broad-spectrum incandescent and fluo-
rescent lights. As LED arrays are highly malleable with
regard to spectral composition, they can be tailored to
reduce or increase insect catches depending on the need.
Wavelength
In comparisons of lamps emitting narrow wavelengths
of light, mosquitoes, Culicoides midges and sand flies
have generally been found to be attracted in higher
numbers by short wavelengths, such as UV, blue (450–
495nm) and green (495–570nm) light [17–19, 46, 50,
52, 58–70]. A UV light trap emitting predominantly
at 325 nm caught fewer mosquitoes than UV traps
emitting at 350–365nm [50], possibly suggesting that
shorter UV wavelengths are less attractive to mosqui-
toes than longer UV wavelengths. Yellow wavelengths
(570–590 nm) can be either attractive or unattrac-
tive [15, 17]. Species vary with regard to which wave-
lengths they are biased towards (See Additional file1
for synthesis).
Longer wavelengths are usually less attractive to bit-
ing flies [15, 17, 18, 58, 62, 66–68, 70, 71]. However,
a few studies have reported catching more Phleboto-
mine sand flies with red wavelengths (620–750 nm)
than with lights emitting shorter wavelengths [46, 72,
73]. Sand flies are anautogenous, meaning they pri-
marily feed on sugar and only require a blood meal
to produce viable eggs. An attraction to longer wave-
lengths may help sand flies locate food plants [73],
although this theory fails to explain why mosquitoes
and midges, which are also anautogenous, do not seem
to share this red attraction. In one study, resting boxes
illuminated with red or infrared wavelengths caught
more mosquitoes than boxes emitting shorter wave-
lengths [66], raising the possibility that red-attracted
sand flies were seeking a resting place. Further studies
are necessary to determine why high numbers of sand
flies were caught using long wavelength LEDs. High
catches with short wavelength light and small catches
with long wavelength light are consistent with fly
spectral sensitivities (Fig.1), suggesting that spectral
Fig. 2 Spectral distribution of three light types: incandescent, compact fluorescent (CFL) and neutral-white, light-emitting diode (LED). Figure
adapted from [116]
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Wilsonetal. Parasites Vectors (2021) 14:28
sensitivities can be used to predict wavelength attrac-
tiveness in some taxa.
Colour vision has not been conclusively shown in the
Culicidae, Ceratopogonidae or Phlebotominae. How-
ever, a few studies have examined the attractiveness of
a wavelength over a range of intensities. For Culicoides
brevitarsis, catches were higher for the green (520 nm)
wavelength than for broad-spectrum incandescent light
even when the green light was half the intensity of the
incandescent [18]. Catches of Culicoides sonorensis
were higher under UV (395 nm) light than under blue
(460nm) light, despite the latter being twice as intense
[70]. Similarly, Lutzomyia longipalpis catches were higher
under UV (350nm) and blue/green (490–546 nm) light
than under a violet (400nm) control light, regardless of
whether the former were lower, equal or higher in inten-
sity than the control [64]. Further studies are needed to
confirm the existence of colour vision in these species.
Wavelength discrimination independent of intensity has
not yet been demonstrated in a mosquito.
Intensity
Traps using more powerful lights tend to catch more
mosquitoes, midges and sand flies than those using dim-
mer ones [46, 49, 71, 74–76], and increasing the intensity
of a given wavelength will generally increase the attrac-
tiveness of the light [18, 76–78]. Two studies have sug-
gested an upper threshold of intensity above which biting
fly catches either reach an asymptote [18] or begin to
decrease [77]—although such a threshold requires cor-
roboration. It is likely that upper threshold varies by taxa
and wavelength. Recording thresholds would ensure
energy is not wasted by increasing intensity beyond that
where insect catches no longer increase, as well as pre-
venting unnecessary light pollution.
Studies into the attractiveness of specific wavelengths
often do not control for light intensity. When ‘green’
and ‘blue’ LEDs are used, the ‘green’ almost always has a
higher luminous intensity—the quantity of visible light
emitted by a source at a given angle—than the ‘blue’ [15,
59, 60, 61, 62, 66, 69, 72, 73]. is is because LED bright-
ness is standardised against the human eye. Two lights
that appear equally bright to humans may be noticeably
different to other animals. e larger catches around
LEDs emitting green wavelengths may therefore be a
result of the greater luminous intensity of these LEDs,
and not a result of the wavelength. Complicating matters
is that the attractiveness of one wavelength over another
is influenced by whether both lights are at a low, medium,
or high intensity. In one study, a blue (470 nm) LED
attracted a higher number of Anopheles mosquitoes than
a green (520nm) LED of equal luminous intensity [76],
but increasing the intensity of the green LED had a larger
effect on Anopheles catches than increasing the intensity
of the blue. Similarly, a blue (470nm) LED was found to
be more attractive than the equivalent green (520 nm)
LED to sand flies, yet increasing luminous intensity sig-
nificantly increased sand fly catches with the green LED
but not the blue LED [78]. Finally, in choice-chamber
experiments, when all the lights were at a low inten-
sity, slightly more Lutzomyia longipalpis sand flies were
attracted to blue–green (490–546nm) wavelengths than
they were the UV (350nm) wavelength. However, when
all the lights were at a higher intensity, more sand flies
were attracted to UV light than the blue/green light [64].
is interaction between intensity and wavelength occurs
because the different photoreceptor classes have differing
sensitivities to light [64].
As users of domestic lights and conductors of vector
surveys will use a variety of light intensities—due to cost,
application and availability limiting the strength of the
power supply—understanding the relationship between
wavelength and intensity will allow for the most appro-
priate wavelength to be chosen for a given intensity.
Contrast
As nocturnal insects have poor visual resolution, contrast
with the background is an important component of visual
attraction in host location and in flight [42, 79]. Studies
on the visual attraction of biting flies have shown that
the attractiveness of an object can be influenced by sur-
rounding vegetation and ambient lighting. Green cloth
was found to be less attractive than its spectral reflec-
tivity would suggest when used against a green, spruce
background [80]. Conversely, red cloth was more attrac-
tive when against that same background. In a study on
the colour preferences of the mosquito Mansonia pertur-
bans, white-coloured traps were unattractive during the
day but highly attractive at night. e reverse was seen
for the blue-coloured trap [81].
ere has been little research into how environment
affects the conspicuousness of emitted light (direct from
a light source) as opposed to reflected light (colour of an
object). Insects may behave differently towards emitted
light and reflected light of the same ‘colour’. For example,
red objects are reported to be attractive to mosquitoes
[80–82], whereas red light is not (see section Wave-
length). Understanding how the environment affects the
visibility of certain lights could help to explain conflict-
ing results for the same species [60, 61]. It could also
potentially identify the most conspicuous wavelength of
light for a given environment for vector surveillance and
inform homeowners which colour backgrounds increase/
decrease the attractiveness of white domestic lighting.
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Wilsonetal. Parasites Vectors (2021) 14:28
Competing light sources
It is well established that light traps decrease in effective-
ness as ambient light levels increase. When using light
traps, it is sometimes recommended that sampling not
take place on nights during or around a full moon due to
the reduced catches on these nights [62, 83–89]. ese
reduced catches are unlikely to be caused by low insect
flight activity. Mosquito flight activity has been shown to
be higher during the full moon when trapping with truck,
suction and animal-baited traps [90, 91]. Lunar phase had
a significant effect on light trap catches, but not on sticky
trap catches [89]. e reduced ability of the light trap to
collect insects may be due to competition between light
from the moon and that from the light trap [92]. Further-
more, light traps are not considered a suitable monitor-
ing tool in northern latitudes where light levels do not
fall below twilight, or in areas with significant light pollu-
tion [93–95]. High levels of background illumination may
reduce the contrast and, therefore, attractiveness of the
trap [96].
Vector surveillance may become more difficult as light
pollution increasingly pervades rural areas where vec-
tors are endemic [97]. However, background illumina-
tion appears to have a stronger effect on some lights
more than others. Moonlight reduced catches more with
incandescent light than with green (520nm) LEDs [62].
Researchers may be able to mitigate the impact high
ambient lighting, from moonlight or light pollution, has
on vector surveillance by using LEDs emitting certain
wavelengths.
Range ofattraction
e ‘range of attraction’ is defined as the maximum dis-
tance from which a light begins to attract insects. Knowl-
edge of attraction ranges is used to determine how far
apart light traps must be for them to not influence each
other during sampling. Light traps with larger attrac-
tion ranges are able to sample insects from a wider area,
which increases catch sizes and reduces the number of
traps needed to sample a given area. Domestic lights with
large attraction ranges would potentially attract a higher
number of vectors. Attraction range is likely influenced
by many factors, such as light intensity, bulb type, host
presence, environment and study species [10, 98, 99].
For Lutzomyia sand flies, the range of attraction of an
incandescent light trap has been estimated to be between
2 and 6 m [100, 101], and for Anopheles mosquitoes,
<5m [102]. Studies on Culicoides midges, however, have
produced highly variable results. e range of attraction
of a Centres for Disease Control and Prevention (CDC)
UV-light trap was approximately 15m [103], and for the
Onderstepoort black-light trap, the range has been esti-
mated as ~1m [99], ~3m [10] or as high as ~30m [98].
e studies with the large attraction ranges [98, 103]
did not have livestock in the general vicinity of the light
traps, whereas the studies with small attraction ranges
[10, 99] did. us, additional olfactory cues may have
caused the considerably shorter attraction ranges. Flies
may have been primarily attracted to the animal and only
responded to the light traps when at close proximity.
Flicker
e critical fusion frequency (CFF) is the frequency at
which a flickering light becomes indistinguishable from
a continuous light source. Human eyes have a CFF of
50–60Hz, whereas diurnal flying insects, including the
tsetse fly, have CFF of > 100Hz [104]. Nocturnal insects
have an average CFF of 70Hz, although the average for
nocturnal, flying insects is likely to be higher due to the
visual demands of flight [57]. e higher CFFs of insects
imply they can perceive flickering that humans cannot,
and this may affect light attraction.
Very few studies have examined fly behaviour towards
flickering lights. For white fluorescent lights, the mos-
quito, Culex quinquefasciatus, and the housefly, Musca
domestica, were found to be more attracted to the direct
current (DC)-powered, non-flickering light than the
alternating current (AC)-powered, flickering light [105].
In another study, fewer Diptera were also caught with
white LEDs flickering at 120 Hz than with LEDs with a
constant light output [57]. Finally, in two choice experi-
ments between flickering and non-flickering white fluo-
rescent lamps, lamps flickering at 10 and 4Hz were less
attractive to M. domestica than the 40,000 Hz control
[106]. ere was no difference in terms of attraction
between the control light and light frequencies > 10Hz.
e author of the study noted that the 10 and 4Hz lights
caused flies to exhibit an “escape response” towards the
non-flickering lamp and suggested that the sudden
reduction in light intensity mimics an attack from a pred-
ator. However, in another study on UV fluorescent lights,
the 100 Hz flickering light caught more M. domestica
than a DC-powered, non-flickering light [107]. In that
study, the flickering light was more attractive even at half
the intensity of the non-flickering light. Whether flicker
is considered attractive may be influenced by the spec-
tral composition of the light: a flickering UV light may be
attractive whereas a flickering white light may not.
e effects of flicker on biting flies require further
investigation. Domestic lights traditionally operate on
AC, where the current alternates on and off. LEDs react
to these current changes much quicker than incandes-
cent and fluorescent lights, resulting in a more pro-
nounced flicker. If flickering lights are less attractive,
then this is an added benefit to using LEDs for external
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Wilsonetal. Parasites Vectors (2021) 14:28
lighting. Changes to the flicker frequencies of domestic
lights could also allow for lights to be made less attractive
without impoverishing the colour rendering.
Conclusions
To improve biting fly capture rates, it is recommended
to use high-intensity, short wavelength LEDs; and to trap
in areas with no competing light sources. Green light
has the advantage over the similarly attractive UV light,
as the catches are ‘cleaner’—collecting fewer non-target
insects, like Lepidoptera [19].
A minimally attractive light source would be a dim, red
LED. However, the fewer the wavelengths composing a
light source, the poorer the colour rendering—defined
as the ability of a light to accurately represent the colours
of objects [57]. With lights intended for home illumina-
tion, it is important to strike a balance between colour
rendering and attractiveness to insects. It is also impor-
tant to keep in mind that conditions such as colour blind-
ness may exacerbate any colour rendering issues the light
has. White light can be created by combining three or
two narrow wavelengths (e.g. red–green–blue, blue–yel-
low, red–cyan). ese lights would have improved colour
rendering over a mono-chromatic light. It is unknown
whether biting flies would perceive a white light com-
posed of few, narrow wavelengths as being equally attrac-
tive as a broad-spectrum white light. However, for a light
composed of few wavelengths to be equally bright as a
broad-spectrum light, the intensity of those wavelengths
would need to be higher. Increasing the intensity of cer-
tain wavelengths may counteract the effects of remov-
ing other wavelengths, resulting in a light that is no less
attractive than a typical white light. Another potential
solution is to reduce the strength of short-range wave-
lengths in a broad-spectrum light, thereby creating a
‘warm’ toned white light. is could potentially reduce
the attractiveness of the light whilst keeping the colour
rendering relatively high. However, experiments compar-
ing insect catches between white LEDs of subtly different
spectral emissions have produced conflicting results [54,
56, 108–110], and further research is therefore required.
If the spectral composition of a lamp cannot be
altered, changing other aspects of the light, such as back-
ground contrast, intensity or flicker might still reduce
its attractiveness. For example, homeowners may be
advised which colours to paint their house to reduce the
Fig. 3 Map showing the countries where field studies into wavelength preferences on Culicidae (triangles), Ceratopogonidae (circles), and
Phlebotominae (squares) have been carried out. Map is adapted from QGIS
Content courtesy of Springer Nature, terms of use apply. Rights reserved.

Page 8 of 11
Wilsonetal. Parasites Vectors (2021) 14:28
attractiveness of their outdoor lighting. Homeowners,
particularly those in countries with high rates of vector-
borne disease, may be willing to accept a minor reduc-
tion in light brightness. Any benefits of light flickering
must also be weighed against the health risks, especially
to those with photosensitive epilepsy.
A number of studies have shown that Lutzomyia lon-
gipalpis, Culicoides brevitarsis, and Culicoides sonoren-
sis are more attracted to certain wavelengths over others
regardless of light intensity [18, 64, 70]. ese insects
may possess colour vision. A convincing test for colour
vision was pioneered by von Frisch [111], who trained
honeybees to associate blue or yellow cards with a sugar
solution, and then had the bees choose between the col-
oured card and multiple shades of grey. It is assumed
that if the animal is only using achromatic signals, then
at least one shade of grey would be indistinguishable
from the training colour. rough these experiments, von
Frisch was able to prove that honeybees possess colour
vision. Mosquitoes can also be trained to associate sugar
solution with certain visual cues [27], suggesting similar
colour vision experiments can be performed on flies.
Further studies are needed to corroborate which wave-
lengths are attractive (Additional file 1) and gain data
on more species. A comprehensive list of species-spe-
cific differences would potentially allow for surveys to
be developed for particular taxa. In this review, almost
half of the studies examining the attractiveness of vari-
ous wavelengths took place in the Americas (Fig.3). Few
studies have been carried out in sub-Saharan African
countries, and fewer still in Asian-Pacific countries, even
though these countries would greatly benefit from infor-
mation on vector light attraction. Not only is the burden
of vector-borne disease high in these countries [4], but
the difficultly in acquiring olfactory bait means light trap-
ping is the easiest and most reliable surveillance method
[112].
Behavioural data would also benefit from support-
ing physiological data. e behaviour of sand flies in
response to red light has led to suggestions that this fam-
ily possesses long wavelength receptors [72, 73], although
Mellor etal. [33] found no evidence of a long wavelength
receptor; therefore, further investigation is needed.
Additionally, no sensitivity data have been collected for
the Ceratopogonidae. Culicoides species can potentially
be divided into two groups in terms of light preference,
namely green-attracted and UV-attracted [19], and the
spectral sensitivities may reflect this grouping. Physi-
ological studies on a wider variety of species may reveal
differences between families or between diurnal and noc-
turnal flies.
Understanding how wavelength, intensity, background
contrast, range of attraction and light flicker interact
with each other would provide a more complete picture
of light attraction in biting flies. More research is also
needed on aspects of light attraction not discussed in
this review, such as light height [113, 114], light polari-
sation [115], time of day effects [40] and the presence of
reflective surfaces [72]. Finally, future studies should con-
trol for thermal emissions, due to the attractiveness of IR
light, as well as intensity, as its effects may not be consist-
ent across wavelengths [64, 76, 78].
In summary, in this review we outline how light trap-
ping can be made more efficient and we highlight how,
despite current knowledge of how to reduce insect
attraction to lights, modifying domestic lights remains
as a challenging though potentially important research
direction.
Supplementary Information
The online version contains supplementary material available at https ://doi.
org/10.1186/s1307 1-020-04530 -3.
Additional le1. Wavelength preferences/biases of various species of
Ceratopogonidae, Culicidae, and Phlebotominae in experiments compar-
ing catches between lights of different wavelengths. What constitutes a
preference/bias was determined on a case-by-case basis. Species with
inconclusive preferences/biases are not listed here. Colour is used in cases
where dominant wavelength is not specified.
Abbreviations
CDC: Centres for Disease Control; CFL: Compact fluorescent lamp; CFF: Critical
fusion frequency; IR: Infrared; LED: Light-emitting diode; R: Retinula cell; Rh:
Rhodopsin; UV: Ultraviolet.
Acknowledgements
Not applicable.
Authors’ contributions
RW was involved in the conception, drafting, writing and editing of the
manuscript. AW, NR and GJ were involved in the writing and editing of the
manuscript. All authors read and approved the final manuscript.
Funding
RW was supported by a NERC iCASE PhD studentship partnered with Integral
LED, UK (Grant NE/R008701/1). Funders did not contribute to the conception,
writing or editing of the manuscript or the decision to publish.
Availability of data and materials
The data generated during this study is included in this published article in
Additional file 1.
Ethics approval and consent to participate
Not applicable.
Consent for publication
Not applicable.
Competing interests
RW was supported by a NERC iCASE PhD studentship partnered with the LED
manufacturer Integral LED, UK. Funders did not contribute to the conception,
writing, or editing of the manuscript or the decision to publish.
Content courtesy of Springer Nature, terms of use apply. Rights reserved.

Page 9 of 11
Wilsonetal. Parasites Vectors (2021) 14:28
Received: 8 September 2020 Accepted: 7 December 2020
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