DOI: 10.1111/eea.

12648

The photokinesis of oriental fruit flies, Bactrocera

dorsalis, to LED lights of various wavelengths
Huan Liu1 , Zhou Gao1, Shu-Zhen Deng2, Feng-Qin Cao3 & Yong-Yue Lu1*
1
Department of Entomology, South China Agricultural University, Guangzhou 510642, China, 2Institute of Biotechnology,
Zhejiang University, Hangzhou 310058, China, and 3College of Environment and Plant Protection, Hainan University,
Haikou 570228, China
Accepted: 20 July 2017

Key words: compound eye, methyl eugenol, phototaxis, monitoring trap, fecundity, mortality rates,
light-emitting diode, Diptera, Tephritidae

Abstract The oriental fruit fly, Bactrocera dorsalis Hendel (Diptera: Tephritidae), is an invasive pest of orchards
around the world, particularly in Asian countries such as China. Light traps offer a potential means
for pest monitoring and management. This study aimed to evaluate the sensitivity of the fly to light
and investigate the impact of monochromatic light in the sensitivity spectrum on B. dorsalis. Six light
wavelengths in LEDs – green (522 nm), yellow (596 nm), blue (450 nm), red (633 nm), purple
(440 nm), and white (compound light) – were adapted to test responses of 5-, 10-, and 20-day-old
B. dorsalis adults kept in laboratory conditions. We also tested the effects of green and red lights on
pupal development and adults’ life activities. The results indicated a phototaxis preference rank in
B. dorsalis adults to monochromatic LEDs with, in decreasing order, green, yellow, purple, blue, and
red. Moreover, positive phototaxis significantly increased with age. Male adults are more sensitive
than female adults to test lights, mainly at the age of 10 and 20 days. Emergence rates of pupae
exposed to 12 and 24 h green light daily were 42 and 67%, respectively, whereas controls held in red
light emerged at 33 and 37%, respectively. Furthermore, body weight, female fecundity, and mortal-
ity of B. dorsalis in night-time exposure of green light (from 21:00 to 09:00 hours; during daytime
flies were illuminated by white LED light) were significantly higher than in red-light test groups and
dark controls. In conclusion, B. dorsalis displayed preference toward green light, and fly age and gen-
der seemed to significantly impact the phototactic behavior. Green LED light exposure during night-
time remarkably improved the emergence rates of B. dorsalis, and it enhanced growth, development,
and ovipositing peak period, but decreased adult lifespan. This research lays a foundation for the
development of new trap models, e.g., with green sticky cards or green light, for monitoring and con-
trol of B. dorsalis in the field.

is an important strategy in controlling Tephritidae (Di-

Introduction
Sight plays an essential role in insect behavior – compound
eyes process light stimuli from the environment, leading to
food sources, reproductive partners, oviposition sites,
hosts, prey, or detecting predators from a distance (Hom-
berg & Paech, 2002; Stalleicken et al., 2006). When the
photoreceptor cells of compound eyes receive motiva-
tional light stimuli, insects display positive or negative
movement. Trapping technology based on chromatic cues
*Correspondence: Yong-Yue Lu, Department of Entomology, College
of Agriculture, South China Agriculture University, 483 Wushan
Road, Guangzhou, Guangdong 510642, China.
E-mail: luyongyue@scau.edu.cn
ptera). Females of the olive fruit fly, Bactrocera oleae Gme-
lin, were more attracted to yellow or orange spheres than
to red, blue, green, black, or white spheres (Katsoyannos,
1989). Yellow fruit-mimicking spheres of 7 cm in diame-
ter were the most attractive to Mediterranean fruit fly, Cer-
atitis capitata Wiedemann (Katsoyannos, 1989; Cornelius
et al., 1999). A previous study has shown that a green
stimulus is the most suitable cue for trapping melon fruit
fly, Bactrocera cucurbitae Coquillett (Xue & Wu, 2013). Li
et al. (2017) found that yellowish green and yellow were
the optimum colors for trapping Bactrocera tau Walker on
sticky panels in the field.
The oriental fruit fly, Bactrocera dorsalis Hendel, is a
devastating agricultural pest with health implications in

102 © 2018 The Netherlands Entomological Society Entomologia Experimentalis et Applicata 166: 102–112, 2018

the tropical and subtropical countries. Bactrocera dorsalis
is highly polyphagous with multiple overlapping genera-
tions, which has been reported to damage more than 250
species of fruit and vegetable crops (Li et al., 2012; Shen
et al., 2012). Females of B. dorsalis will lay eggs inside
fruits, wherein larvae will feed until they come out and
drop to hide in the soil for pupation. Therefore, the appli-
cation of chemical control of the hidden immature stages
is difficult and insecticides proved unsuccessful. Moreover,
B. dorsalis field populations developed resistance against
organophosphates, synthetic pyrethroids, and abamectin
insecticides (Hsu & Feng, 2000; Jin et al., 2011; Vontas
et al., 2011). The para-pheromone methyl eugenol (ME)
is widely and effectively used to detect, control, and eradi-
cate male adults of B. dorsalis worldwide (Vargas & Pro-
kopy, 2006; Lin et al., 2012; Pagadala et al., 2012).
However, ME was considered carcinogenic to humans by
the National Toxicology Program of the United States
Department of Health and Human Services, hampering its
long-term use (Miller et al., 1983; Shelly, 1997; Smith
et al., 2002; Zheng et al., 2012). In addition, ME can only
trap sexually mature male flies, but will not affect imma-
ture forms or females. Thus, it is urgent to explore a more
ecologically friendly, simple, and effective method for con-
trolling B. dorsalis.
Bactrocera dorsalis adults’ diurnal host-finding behavior
is strongly affected by visual cues. Therefore, the best strat-
egy to attract and lure oriental fruit flies is by using traps
based on visual stimuli (Prokopy & Owens, 1983; Wu
et al., 2007). By hanging fruit-mimicking spheres of differ-
ent colors in guava (Psidium guajava L.) trees, Vargas et al.
(1991) demonstrated that yellow or white spheres would
be useful devices for monitoring the oriental fruit fly. In
subsequent field tests, Cornelius et al. (1999) found that
yellow spheres captured more female oriental fruit flies
than yellow rectangular blocks, or than spheres of other
colors. Huang & Lin (2004) found that female flies of
B. dorsalis prefer to oviposit into mature yellow fruit in
the field. Unfortunately, these colors were not quantified.
Wu et al. (2007) found that green light stimuli (500–
570 nm) enhanced the attractiveness of B. dorsalis by use
of green paper, whereas blue light stimuli (380–500 nm)
diminished the attractiveness by using electroretinogram
technology. It should be noted that there was considerable
color difference between printed paper sheets. The use of
monochromatic light sources may solve this issue. How-
ever, no research has yet examined the light responses of
B. dorsalis.
A light-emitting diode (LED) is a semiconductor device
that produces highly monochromatic light with little heat,
it has a long lifetime, low energy consumption, and a high
degree of safety (Duehl et al., 2011; Cho & Lee, 2012; Jeon
Evaluation of phototaxis by Bactrocera dorsalis 103
et al., 2012). The solid state physics of LEDs are an advan-
tage for controlling pest behavior and more efficient in
practical application. LED traps have been manufactured
based on insect responses toward the lighting and were
widely used for monitoring and control of pests such as
Culicoides brevitarsis Kieffer (Bishop et al., 2006), Tri-
bolium castaneum Herbst (Duehl et al., 2011), Euscepes
postfasciatus Fairmaire (Katsuki et al., 2012), and Helico-
verpa armigera H€ ubner (Yoon et al., 2012). A variety of
abiotic factors can influence the efficiency of light traps,
including the light wavelength, intensity, time of exposure,
and the spectral composition of the light (Shimoda &
Honda, 2013; Sun et al., 2014). Also, the responsiveness of
insects to light appears to differ between the sexes. Due to
their flying capability (Venter et al., 2009), structure of
compound eyes (Lau & Meyer-Rochow, 2007; Meyer-
Rochow & Lau, 2008), and physiological status (Bourg &
Badia, 1995; Belmain et al., 2000; Castrejon & Rojas,
2010), the photoresponsive behavior of female and male
adults is inconsistent. For instance, males and gravid
females of Indian meal moth, Plodia interpunctella
(H€ubner), exhibited preference to blue light, unlike virgin
females (Cowan & Gries, 2009). Females of Loxostege sticti-
calis (L.) and Anomala corpulenta Motschulsky displayed
stronger phototactic responses than males (Jiang et al.,
2010, 2015). When reaching the age of 12 days, adult
males of Xestobium rufovillosum De Geer displayed a pref-
erence for light, whereas females increasingly preferred the
dark (Belmain et al., 2000). The study of innate phototac-
tic behavior can provide the basis for designing electric
insect killers (Shimoda & Honda, 2013).
The phototropic responses of B. dorsalis males and
females of different ages toward multiple LED wave-
lengths, and the influence of monochromatic lights on
biological parameters have not been studied before. This
work tests the preference of B. dorsalis to six LED wave-
lengths in the laboratory. Our results may aid in develop-
ing sustainable, secure, and more efficient light traps for
monitoring B. dorsalis.
Materials and methods
Insects
The B. dorsalis strain was maintained in a laboratory in
South China Agricultural University, at 27
1 °C,
75
1% relative humidity, and L14:D10 photoperiod.
Hatched larvae were maintained on an artificial diet
including sugar (9%), yeast (15.1%), nipagen (0.15%),
sodium benzoate (0.15%), wheat germ oil (0.15%), citric
acid (1.7%), and water (73.8%) (Chang et al., 2006). Adult
flies were reared in cages and fed another artificial diet
consisting of 1:1 yeast extract:dry sugar (wt/wt).
104 Liu et al.

Materials
Based on previous research on phototaxis in Tephritidae
(Wu et al., 2007; Xue & Wu, 2013; Li et al., 2017), six LED
light colors (green, red, yellow, blue, purple, and white;
Table 1) were selected and customized in YuanEr Tech-
nology, Shenzhen, China (Sun et al., 2014). Sartorius-
BP121S type electronic analytical balance was purchased
from Sartorius AG. The Electronic timing socket was pur-
chased from Bull International Electrical Group.
Phototaxis device
In view of the strong flight and climbing ability of
B. dorsalis adults, we designed a phototactic test device
(made by Guangzhou Qianhui Chemical Glass Instru-
ment, Tianhe, Guangzhou, Guangdong, China) (Fig-
ure 1). Each device (25 9 25 9 30 cm) consisted of
transparent acrylic to allow for light exposure, compris-
ing three chambers: a phototactic response chamber,
an activity chamber, and a dark chamber (Figure 1B–
D). Chambers were separated by an opaque partition
with nine insect entrance holes (2 cm diameter). The
LEDs were connected to a rheostat to adjust light
intensities. The light intensity of LEDs was measured
using a CL-200F spectrum illuminometer (Kexing Pho-
toelectric, Hangzhou, China) at the center of the
chamber (30 cm from the light source). The precise
spectral characteristics of LED lights were measured by
a CL-200FZ photoelectric properties integrated test sys-
tem, which consists of a CL-200F spectrum illumi-
nometer, an integrating sphere, and system software
(Kexing Photoelectric).
Phototactic response
The phototactic responses of oriental fruit flies were exam-
ined under set conditions of light wavelength and fly age.
When oriental fruit flies were 5, 10, or 20 days of age, 100
flies with a 1:1 male:female ratio were chosen as test sub-
jects. Prior to each bioassay, we held the flies in the dark
for 30 min, and then placed them in an activity chamber
Figure 1 Device used for testing phototaxis behaviour of
Bactrocera dorsalis adults. (A) Light-emitting diode (LED) light
source; (B) phototactic response chamber; (C) activity chamber;
(D) dark chamber; (E) insect entrance holes; (F) opaque barrier.
(Figure 1C). All experiments were carried out inside a sin-
gle dark room during 17:00 and 20:00 hours. Exposure to
LED light lasted 2 h and light intensity was 200 lx. The
control groups were tested in complete darkness. The
response of B. dorsalis to LEDs was estimated by counting
the males and females inside each test chamber. The pho-
totaxis ratios of females and males were calculated using
the following formula:
Phototaxis rate ð%Þ ¼ ½ðno. flies in chamber B in
treatment group  no. flies in chamber B in control
groupÞ=no. test flies  100%;
where chamber B indicates the phototactic response
chamber (Figure 1B). In total, each experiment was repli-
cated 59. Each fly was used only once. The testing devices
were washed with 75% alcohol after each test, and left to
dry indoors for at least 2 h to remove odors and traces left
by previous insects. During experiments, the room was
maintained at 27
1 °C and 75
1% r.h.
Effect of green and red lights on pupa emergence
Pupae formed within 24 h were separated haphazardly
into six groups of 50, each put into a 100-ml beaker. The
beakers were placed inside corresponding dark boxes
(30 9 30 9 30 cm). The flies were exposed to green or
red LED light (at 200 lx) for 0, 12, or 24 h every day. We

Table 1 Spectral characteristics of light-

Peak Wavelength Illuminance Irradiance Radiant
emitting diode (LED) colors
Color wavelength (nm) range (nm) (lx) (W m2) flux (W)
Green 522 480–580 337.70 0.47 0.48
Red 633 580–670 279.63 0.63 0.64
Yellow 596 540–620 298.68 0.48 0.51
Blue 450 420–500 225.14 0.82 0.86
Purple 440 400–480 219.70 0.53 0.53
White 452 (main), Broad 327.50 0.60 0.63
570 (secondary) spectrum

The LED lights spectral parameters test distance is 30 cm.


counted the emerged adults daily at 10:00, 15:00, and
20:00 hours until all adults emerged. Then we calculated
the pupal emergence rate. All treatments were repeated
39.
Influence of nighttime exposure to green or red lights on adult
weight, mortality, and female fecundity
Groups of 200 newly emerged adults (within 24 h) with a
1:1 male:female ratio were selected haphazardly and placed
in a cage (35 9 35 9 35 cm). Treatment groups were
then illuminated with green or red LED lights (200 lx)
daily, from 21:00 to 09:00 hours. Control groups were
kept in the dark at the same time. From 09:00 to
21:00 hours, both experimental and control groups were
illuminated by white LED light (200 lx; Figure 2). All flies
were supplied the same food and water daily. Each treat-
ment was repeated 39. Individual adult weight was mea-
sured every 5 days. We recorded the number of dead flies
at 10:00 hours every day until all flies were dead. Thus,
adult weight and mortality rates for each treatment and
control were calculated.
When tested flies reached sexual maturity (15 days of
age), we calculated the female fecundity every 3 days. A
plastic centrifuge tube (50 ml), with small oviposition
holes punctured into it using a 19 gauge syringe needle,
was used as an artificial egging device. Subsequently, we
poured 1 ml self-made orange juice (as an oviposition
stimulus) into the egging devices evenly, placed them in
the flies’ cage at 15:00 hours, and allowed oviposition for
4 h. Immediately after the 4 h we counted the eggs and
calculated the fecundity of female adults. Each experiment
was repeated 39.
Figure 2 Device used for testing the effect
of green and red light nighttime
illumination on Bactrocera dorsalis adults.
From 09:00 to 21:00 hours, treatment and
control groups were illuminated by white
LED light (200 lx). Treatment groups were
illuminated daily with 200-lx green or red
LED lights from 21:00 to 09:00 hours.
Control groups were kept in the dark at the
same time. [Colour figure can be viewed at
wileyonlinelibrary.com]
Evaluation of phototaxis by Bactrocera dorsalis 105
Statistical analysis
All analyses were carried out with SAS v.9.20 software
(SAS Institute, Cary, NC, USA). Datasets of adult photo-
tactic response rates to various LED wavelengths, weight,
female fecundity, and mortality were checked for normal-
ity of distribution and homogeneity of variances with
Shapiro–Wilk and Levene’s tests, respectively. If data were
normally distributed and had similar variances, then
means were compared by one-way ANOVA. Significant
ANOVA results were followed by mean separation by
Duncan’s multiple range test (a = 0.05). Non-normally
distributed data were analyzed with a nonparametric
Kruskal–Wallis test to compare medians (a = 0.05), fol-
lowed by a Mann–Whitney test for pairwise comparisons,
if applicable. The attractiveness of purple and blue light to
flies, color preferences between female and male flies, and
pupal emergence rates between green and red light treat-
ment groups were compared with independent-samples t-
test (a = 0.05). Results were plotted with Origin v.7.5
(OriginLab, Northampton, MA, USA).
Results
Phototaxis to different wavelengths of light
In control groups, the mean (
SEM) number of 5-, 10-,
and 20-day-old B. dorsalis adults in the phototactic
response chamber (Figure 1B) was 6.0
0.84, 6.6

1.12, and 6.2
0.80, respectively – much lower than that
in the activity chamber (Figure 1C), which was 89.0

1.10 (F2,8 = 2 324.33), 87.6
2.06 (F2,8 = 643.97), and
86.0
1.22 (F2,8 = 1 351.28, all P<0.0001), respectively
(Figure 3). After 30 min of conditioning in the dark, 5-,
106 Liu et al.

10-, and 20-day-old adults displayed clear phototaxis

toward all tested monochromatic and white lights, with
the strongest behavioral response toward green, followed
by white, then yellow. Green light attraction was signifi-
cantly different to all other monochromatic lights (Fig-
ure 3). The white (composite) light also was strongly
attractive to flies, and red light was the least attractive, with
most flies staying in the activity chamber. There was no
difference in attractiveness of purple and blue light to 10-
(t = 1.72, d.f. = 4, P = 0.16) and 20-day-old flies
(t = 1.19, d.f. = 4, P = 0.30).

Phototaxis at different post-emergence ages

Phototaxis increased with age, with newly emerged adults
presenting lowest phototaxis (Figure 4). The mean
(
SEM) phototactic responses of 10- and 20-day-old
adults to green light were 48.2
0.86 and 54.4
1.60%,
respectively – significantly higher than in 5-day-old adults
(40.4
1.60%) (F2,8 = 20.97, P = 0.0007). Phototactic
responses of 20-day-old adults to white, yellow, purple,
and blue lights were much higher than those of 5-day-old
flies.

Phototaxis of males and females

Overall performance indicated that the number of males
in the phototactic response chamber is higher than that of
females, suggesting clear sexual differences (Figure 5). At
10 and 20 days old, males responded significantly stron-
ger than females to green, white, and yellow light. The
phototactic response of females to red light was stronger
than that of males, significant differences were found in
5- (t = 4.63, d.f. = 4, P = 0.0098) and 10-day-old adults
(t = 4.71, d.f. = 4, P = 0.0093). Although a tendency
toward increased phototaxis reaction of older males is
suggested, the difference is not significant (t = 2.36,
d.f. = 4, P = 0.078).

Influence of green and red light on pupa emergence

Emergence rates under green light exposure for 12 and
24 h daily were 21% (t = 13, d.f. = 2, P = 0.0059) and
45% (t = 7.21, d.f. = 2, P = 0.019) higher than under red
light treatment, demonstrating that green light can signifi-
cantly increase pupal emergence rates (Figure 6). More-
over, following the extending green light irradiation time
every day, the emergence rates increased notably
(F2,4 = 61.30, P = 0.0010).
Influence of nighttime green and red light on fly weight and female
fecundity
Nighttime exposure to green or red light resulted in
increased adult body weight (Figure 7). After 5 nights of
Figure 3 Distribution of Bactrocera dorsalis adult flies (mean
number
SEM) of (A) 5, (B) 10, or (C) 20 days old after
exposure to various light-emitting diode (LED) colors (see
Table 1 for wavelengths). Flies are released in the middle
(activity) chamber (see Figure 1 for setup) and after 2 h
exposure their position is monitored: they are in the
phototactic response chamber (white bars), activity chamber
(light gray bars), or dark chamber (dark gray bars). CK
(control) was in complete darkness. Means within a panel
and within a single LED color capped with different letters
are significantly different (Duncan’s multiple range test:
P<0.05).
 Evaluation of phototaxis by Bactrocera dorsalis 107

Figure 4 Mean (
SEM) phototaxis rate (%) of Bactrocera

dorsalis adults of 5 (white bars), 10 (light gray bars), or 20 days
old (dark gray bars) exposed to various light-emitting diode
(LED) colors (see Table 1 for wavelengths). Means within a
single LED color capped with different letters are significantly
different (Duncan’s multiple range test: P<0.05).

exposure, the mean (
SEM) fly weights did not differ

[green, 7.6
0.09 mg; red, 7.3
0.09 mg; darkness
(control), 7.2
0.03 mg; F2,4 = 4.16, P = 0.11]. After
10–25 nights treatment, flies exposed to green light were
significantly heavier than flies exposed to red or no light –
after 25 nights exposure, flies exposed to green light
weighed 12.3
0.24 mg, those exposed to red light
11.8
012 mg, and the control flies weighed 11.2

0.10 mg (F2,4 = 30.88, P = 0.0037). These results
demonstrate that both green and red nighttime irradi-
ation have a positive impact on the growth and devel-
opment of B. dorsalis adults, whereas the promotion
of green exceeds that of red light.
Green light promoted oviposition by female B. dorsalis
(Figure 8). In the green light treatment groups, oviposi-
tion quickly increased after sexual maturation and peaked
around the 21st day. In the red light test groups and in the
controls, females took 27 days to peak oviposition after
sexual maturation. The highest oviposition rate in the
green light groups was 20.8
0.59 eggs per female, which
is significantly higher than the highest oviposition rate in
the red light (16.1
1.08) and control groups Figure 5 Mean (
SEM) number of male (gray bars) and
female (white bars) Bactrocera dorsalis adult flies of (A) 5,
(14.2
1.63) (F2,4 = 11.76, P = 0.021).
(B) 10, or (C) 20 days old after exposure to various light-
emitting diode (LED) colors (see Table 1 for wavelengths).
Influence of nighttime green and red light on adult mortality rate
Flies are released in the middle (activity) chamber (see
Nighttime exposure to green and red light decreased sur- Figure 1 for setup) and after 2 h exposure their position is
vival of B. dorsalis adults (Figure 9). Mortality of the two monitored in the phototactic response chamber. CK
treated groups and the control group increased monotoni- (control) was in complete darkness. Asterisks indicate a
cally over time; the mortality rate (%) of flies exposed to significant difference between the sexes (t-test: P<0.05).
108 Liu et al.
Figure 6 Mean (
SEM) pupal emergence rate (%) of Bactrocera
dorsalis after exposure for 0, 12, or 24 h per day to green (light
grey bars) and red (dark grey bars) LED lights. Means within a
single LED color capped with different letters are significantly
different (Duncan’s multiple range test: P<0.05); asterisks
indicate a significant difference between light colors (t-test:
P<0.05).
Figure 7 Effect of nighttime exposure for 5–40 days to green and
red LED light on mean (
SEM) weight (mg) of Bactrocera
dorsalis adults. Control was in complete darkness during
nighttime. Means within a single treatment time marked with
different letters are significantly different (Duncan’s multiple
range test: P<0.05).
green light was significantly higher than that of control
flies from day 40 onwards, whereas the mortality rate of
flies exposed to red light was significantly higher than that
of the controls from day 80 onwards. After 80 days of
treatment, mortality in the green and red light treatment
Figure 8 Effect of nighttime exposure for 18–36 days to green
and red LED lights on the fecundity of Bactrocera dorsalis
(mean
SEM no. eggs laid per female). See Figure 2 for setup.
Control was in complete darkness during nighttime. Means
within a single treatment time marked with different letters are
significantly different (Duncan’s multiple range test: P<0.05).
Figure 9 Effect of nighttime exposure for 10–150 days to green
and red LED lights on the mean (
SEM) mortality rate (%) of
Bactrocera dorsalis adults. See Figure 2 for setup. Control was in
complete darkness during nighttime. Means within a single
treatment time capped with different letters are significantly
different (Duncan’s multiple range test: P<0.05).
groups had increased to 18.8
0.60 and 15.7
0.33%,
respectively, and mortality in the control group had
increased to 13.2
1.09% (F2,4 = 20.99, P = 0.0076).
After 150 days of treatment, all insects exposed to green
light were dead, yet 19.8% of flies exposed to red light
and 27.2% control flies were still alive (F2,4 = 223.96,
P = 0.0001). Apparently nighttime exposure to green light

decreases the lifespan of test flies more than red light
treatment.
Discussion
Screening the sensitivity spectrum of insects to light
can aid in controlling insect hazards and epidemics,
e.g., the spread of arboviruses (Antignus, 2000). Most
insects have highly conserved visual pigments, with sen-
sitive spectral absorption between 350 and 700 nm, and
they can perceive and respond to light in this range
(Land, 1997). Insects differ in their sensitivity for speci-
fic wavelengths (Sun et al., 2014). In this study, we
found both sexes in B. dorsalis have a wavelength pref-
erence, especially for green light, followed by white and
yellow, and they are least sensitive to red light. This is
in agreement with previous findings by Wu et al.
(2007) who observed that B. dorsalis adults are more
attracted to green and yellow plates than to red ones,
and by Liang et al. (2016) who observed B. dorsalis
adults have preference for green food items and are less
interested in red food items.
Light irradiation was the required for adult B. dorsalis
foraging, flight, and courtship (Liu & Ye, 2006). Our
observations are consistent with the finding that B. dor-
salis adults were most active in the day, and kept still at
night (or in a dark room). Day-active phytophagous
insects usually display the strongest phototactic reaction
to wavelengths from 500 to 580 nm, while being generally
blind to red (649 nm) and infrared light (Cohnstaedt
et al., 2008; Chen et al., 2013). Insects indiscriminately
respond to green objects including non-host plants (Couty
et al., 2006). Wavelengths preferred by B. dorsalis adults
would correspond to visible green hues. A similar ten-
dency was observed with B. cucurbitae and B. tau (Xue &
Wu, 2013; Li et al., 2017). The green-positive effect could
be attributed to the green sensitivity of the 510-nm pho-
toreceptor of B. dorsalis, which has been described as R8
cells based on the evidence from intracellular electrophysi-
ological recordings (Wu et al., 2007). In nature the surface
color of leaves, flowers, and ripe fruits of the host plants
lays within the range of this sensitive short-wavelength
spectrum – this may partly explain the attraction of green
and yellow lights to tephritids (Prokopy & Owens, 1983;
Drew et al., 2003; Blackmer et al., 2008; Yu, 2011; Tang
et al., 2015). Based on field investigations, both Vargas
et al. (1991) and Cornelius et al. (1999) demonstrated
that yellow fruit-mimicking spheres were strong lures to
B. dorsalis. Yang et al. (2011) studied the color preferences
of B. dorsalis with traps containing a sweet bait hanging
with differently colored balloons in a carambola garden.
They found that the trap performed best where purple
Evaluation of phototaxis by Bactrocera dorsalis 109
balloons were used. These findings do not seem to match
our results. One reason may be that our research was con-
ducted indoors in controlled conditions, where less inter-
ference takes place. Also, the color, shape, and size of traps
are known to affect the response of oriental fruit flies. Dif-
ferences in age, sex, and developmental status of their
stock B. dorsalis may have been another reason for the
inconsistent experimental result.
Age-related phototactic behavioral changes occur in
B. dorsalis adults. The phototaxis rates of 20-day-old flies
were higher than those of 5- and 10-day-old flies. This
finding confirms the conclusions with other insects (Wei
et al., 2000; Jiang et al., 2010; Zhu et al., 2014). Under
various optical environments, the movements of blocking
pigment granules in the pigment cells of insect com-
pound eyes were closely related to the sensitivity to
monochromatic light of different wavelengths and the
status changes of compound eyes (Lau & Meyer-Rochow,
2007; Meyer-Rochow & Lau, 2008). This was a physiolog-
ical basis for the specific sensitivity spectra of different
insect species (Yu, 2011). When illuminating insects with
dark compound eyes, blocking pigment granules inside
pigment cells around the retinal cells move from the end
of cornea to the basilar membrane. They surround the
rhabdomere of each retinal cell. Before reaching the rhab-
domere, the inciting light must go through dark pigment
granules which absorb UV and visible light thus influenc-
ing spectral sensitivity of retinal cells (Chen et al., 1984).
The phototactic rate of B. dorsalis increased with age
probably because compound eyes may not yet be fully
developed in the early imaginal state, when sensitivity of
the blocking pigment cells may be weak, particularly for
short wavelengths such as in the blue and red spectra.
Also, age may affect phototactic tendencies due to a dif-
ference in the level of locomotor activity of flies (Elens,
1972). Males and females display higher flight activity
and are generally more active with the increase in age
(Cui et al., 2016). Mature male and female adults are sen-
sitive to wavelengths as orientation cues during the search
for food, reproductive partners and oviposition sites
(Castrejon & Rojas, 2010).
The phototactic responses of B. dorsalis differed
between the two sexes, with males being more sensitive
to green, yellow, and white lights than females. Photo-
taxis differences between the sexes occur widely among
insects (Meyer, 1978). Many studies reported a greater
abundance of male moths compared to females in light
trap catches (Kiss et al., 2003; Altermatt et al., 2009).
The attractiveness of a blue-LED trap was higher to
males of phlebotomine sand flies than to females during
twilight (Silva et al., 2016). Male adults of Grapholita
molesta Busck have a stronger phototactic reaction to
110 Liu et al.
blue and green light than female adults (Sun et al.,
2014). Males of Carposina sasakii Matsumura preferred
light at 333 nm, whereas females preferred 350 and
405 nm (Hou et al., 1994). This may be linked to gender
differences in flight capability, body physiology, mating
status, and compound eye structure, or to outside condi-
tions such as light sources (Cowan & Gries, 2009; Cheng
et al., 2011; Sun et al., 2014).
Insects are known to have a variety of biological and
physiological responses to various wavelengths of the light
spectrum, including attraction, repulsion, light adapta-
tion, circadian rhythms, light toxicity, photoperiodicity,
and others (Shimoda & Honda, 2013; Tariq et al., 2015,
2017). Circadian rhythms are daily behavioral rhythms of,
e.g., locomotion, flight, feeding, and mating. During the
night, after several minutes or a certain duration of expo-
sure to artificial light, the timing of the diurnal/nocturnal
behaviors of insects may shift and they may exhibit typical
daytime behavior (Walcott, 1969; Okada et al., 1991).
Some diurnal insects have been investigated that are active
in crepuscular or nocturnal light with inherently diverse
irradiance spectra, such as some bees and wasps (Warrant
et al., 2004; Theobald et al., 2007). During our experi-
ment, we observed that behaviors such as foraging, flight,
and mating were activated in B. dorsalis adults when
exposed to green and red lights in the night. As a result, we
found that nighttime exposure to green light promoted
B. dorsalis adult development, accelerating ovary develop-
ment and maturation in a way that hastened the peak per-
iod of oviposition and increased fecundity. Zhou et al.
(1995) pointed out that exposure to medium and long
light wavelengths seemed beneficial to reproduction in
B. dorsalis, as observed in our study. After exposure to
green light the average lifespan of B. dorsalis adults was
found to be decreased. This finding was similar to observa-
tions with Plutella xylostella L., G. molesta, and Tetrany-
chus urticae Koch (Duan et al., 2010; Ismail et al., 2011;
Yu, 2011).
Light traps are based on insect phototactic behavior,
which has been considered as a ‘clean’ form of pest moni-
toring and control (Johansen et al., 2011). Some diurnal
insects can be attracted to light sources at night (Shimoda
& Honda, 2013). In addition, colored devices, such as yel-
low pan traps or yellow sticky traps, are widely used to
capture diurnal pests including aphids, whiteflies, thrips,
and leafminer flies (Mainali & Lim, 2010; Shimoda &
Honda, 2013). Currently, various yellow sticky plates are
widely employed in monitoring and control of B. dorsalis
(Li et al., 2017). We found that its phototactic sensitivity
to green light (522 nm) was greater than to yellow light.
From a pest management perspective, our findings indi-
cate that a stimulus in the green spectra may hold higher
attractive potential to B. dorsalis, thus tests with green
sticky traps or light traps are warranted.
Acknowledgements
We thank Eduardo Fox and Waqar Jaleel for linguistic
correction. We are grateful to the members of our labora-
tory for their cooperation in oriental fruit fly rearing and
treatment. This research was supported by grants from
Guangdong Province College High-Quality Professional
Foundation (No. 246) and National key research and
development project of China (No. 2016YFC1201204).
References
Altermatt F, Baumeyer A & Ebert D (2009) Experimental evi-
dence for male biased flight-to-light behavior in two moth spe-
cies. Entomologia Experimentalis et Applicata 130: 259–265.
Antignus Y (2000) Manipulation of wavelength-dependent beha-
viour of insects: an IPM tool to impede insects and restrict epi-
demics of insect-borne viruses. Virus Research 71: 213–220.
Belmain SR, Simmonds MSJ & Blaney WM (2000) Behavioral
responses of adult deathwatch beetles, Xestobium rufovillosum
de Geer (Coleoptera: Anobiidae), to light and dark. Journal of
Insect Behavior 13: 15–26.
Bishop AL, Bellis GA, McKenzie HJ, Spohr LJ, Worrall RJ et al.
(2006) Light trapping of biting midges Culicoides spp. (Di-
ptera: Ceratopogonidae) with green light-emitting diodes.
Australian Journal of Entomology 45: 202–205.
Blackmer JL, Byers JA & Saona CR (2008) Evaluation of color
traps for monitoring Lygus spp.: design, placement, height,
time of day, and non-target effects. Crop Protection 27: 171–
181.
Bourg EL & Badia J (1995) Decline in photopositive tendencies
with age in Drosophila melanogaster (Diptera: Drosophilidae).
Journal of Insect Behavior 8: 835–845.
Castrejon F & Rojas JC (2010) Behavioral responses of larvae and
adults of Estigmene acrea (Lepidoptera: Arctiidae) to light of
different wavelengths. Florida Entomologist 93: 505–509.
Chang CL, Vargas RI, Caceres C, Jang E & Cho IK (2006) Devel-
opment and assessment of a liquid larval diet for Bactrocera
dorsalis (Diptera: Tephritidae). Annals of the Entomological
Society of America 99: 1191–1198.
Chen DM, Collins JS & Goldsmith TH (1984) The ultraviolet
receptor of bird retinas. Science 225: 337–340.
Chen Y, Luo CW, Kuang RP, Li HW, Chen Z & Liu CY (2013)
Phototactic behavior of the Armand pine bark weevil, Pissodes
punctatus. Journal of Insect Science 13: 1–9.
Cheng WJ, Zheng XL, Wang P, Lei CL & Wang XP (2011)
Sexual difference of insect phototactic behavior and related
affecting factors. Chinese Journal of Applied Ecology 22:
3351–3357.
Cho KS & Lee HS (2012) Visual preference of diamondback
moth, Plutella xylostella, to light-emitting diodes. Journal of
Korean Society for Applied Biological Chemistry 55: 681–684.

Cohnstaedt LW, Gillen JI & Munstermann LE (2008) Light-emit-
ting diode technology improves insect trapping. Journal of the
American Mosquito Control Association 24: 331–334.
Cornelius ML, Duan JJ & Messing RH (1999) Visual stimuli and
the response of female oriental fruit flies (Diptera: Tephritidae)
to fruit-mimicking traps. Journal of Economic Entomology 92:
121–129.
Couty A, Van Emden H, Perry JN, Hardie J, Pickett JA & Wad-
hams LJ (2006) The roles of olfaction and vision in host-plant
finding by the diamondback moth, Plutella xylostella. Physio-
logical Entomology 31: 134–145.
Cowan T & Gries G (2009) Ultraviolet and violet light: attractive
orientation cues for the Indian meal moth, Plodia interpunctella.
Entomologia Experimentalis et Applicata 131: 148–158.
Cui JX, Dong JF, Ren XH, Wu LM, Zuo WQ & Wang Y (2016)
Effects of gender and age (in days) on flight capacity of an
experimental population of the oriental fruit fly, Bactrocera dor-
salis (Hendel). Acta Ecologica Sinica 36: 1292–1302.
Drew RAI, Prokopy RJ & Romig MC (2003) Attraction of fruit
flies of the genus Bactrocera to colored mimics of host fruit.
Entomologia Experimentalis et Applicata 107: 39–45.
Duan Y, Wu RH, Wu YQ, Jiang YL & Zhao MQ (2010) Effects of
LED illumination on the biology of Plutella xylostella. Henan
Agriculture Science 1: 80–82.
Duehl AL, Cohnstaedt AR & Teal P (2011) Evaluating light
attraction to increase trap efficiency for Tribolium castaneum
(Coleoptera: Tenebrionidae). Journal of Economic Entomo-
logy 104: 1430–1435.
Elens A (1972) Influence of aging on behavior of D. melanogaster.
Drosophila Information Services 48: 108.
Homberg U & Paech A (2002) Ultrastructure and orientation of
ommatidia in the dorsal rim area of the locust compound eye.
Arthopod Structure and Development 30: 271–280.
Hou WW, Ma YF, Gao WZ, Li SW & Yang ZJ (1994) A study on
the phototaxis of the peach fruit moth. Acta Entomologica
Sinica 2: 165–170.
Hsu JC & Feng HT (2000) Insecticide susceptibility of the oriental
fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae) in
Taiwan. Chinese Journal of Entomology 20: 109–118.
Huang AF & Lin LF (2004) The harm situation and preventive
measures of Bactrocera dorsalis (Hendel) in Zhangzhou. Fujian
Science and Technology of Tropical Crops 29: 33–34.
Ismail MSM, AboGhalia AH & Soliman MFM (2011) Cer-
tain effects of different spectral colors on some biological
parameters of the two-spotted spider mite, Tetranychus
urticae. Egyptian Journal of Biological Pest Control Papers
3: 27–39.
Jeon JH, Oh MS, Cho KS & Lee HS (2012) Phototactic response
of the rice weevil, Sitophilus oryzae Linnaeus (Coleoptera: Cur-
culionidae), to light-emitting diodes. Journal of Korean Society
for Applied Biological Chemistry 55: 35–39.
Jiang XF, Zhang ZZ & Luo LZ (2010) Phototaxis of the webworm
Loxostege sticticalis to different wavelengths and light intensity.
Plant Protection 36: 69–73.
Jiang YL, Wu YQ, Li T, Gong ZJ, Duan Y et al. (2015) Beha-
vioural responses of Anomala corpulenta Motschulsky
Evaluation of phototaxis by Bactrocera dorsalis 111
(Coleoptera: Scarabaeoidea) to different spectral light. Acta
Entomological Sinica 58: 1146–1150.
Jin T, Zeng L, Lin YY, Lu YY & Liang GW (2011) Insecticide resis-
tance of the oriental fruit fly, Bactrocera dorsalis (Hendel) (Di-
ptera: Tephritidae), in mainland China. Pest Management
Science 67: 370–376.
Johansen NS, V€anninen I, Pinto DM, Nissinen AI & Shipp L
(2011) In the light of new greenhouse technologies. 2. Direct
effects of artificial lighting on arthropods and integrated pest
management in greenhouse crops. Annals of Applied Biology
159: 1–27.
Katsoyannos BI (1989) Response to shape, size and color. Fruit
Flies: Their Biology, Natural Enemies and Control, Vol. 3A
(ed. by AS Robinson & G Hooper), pp. 307–324. Elsevier,
Amsterdam, The Netherlands.
Katsuki M, Omae Y, Okada K, Kamura T, Matsuyama T et al.
(2012) Ultraviolet light-emitting diode (UV LED) trap the
West Indian sweet photo weevil, Euscepes postfasciatus
(Coleoptera: Curculionidae). Applied Entomology and Zoo-
logy 47: 285–290.
Kiss M, Nowinszky L & Puskas J (2003) The ratio of males and
females in the catch. The Handbook of Light Trapping (ed. by
L Nowinszky), pp. 189–194. Savaria University Press, Szom-
bathely, Hungary.
Land MF (1997) Visual acuity in insects. Annual Review of Ento-
mology 42: 147–177.
Lau TF & Meyer-Rochow VB (2007) The compound eye of
Orgyia antiqua (Lepidoptera: Lymantriidae): sexual dimor-
phism and light/dark adaptational changes. European Journal
of Entomology 104: 247–258.
Li XY, Tan JC, Song DB & You LS (2012) Parasitoids of Bactro-
cera dorsalis and their application to bio-control. Biological
Disaster Science 35: 12–17.
Li L, Ma HB, Niu LM, Han DY, Zhang FP et al. (2017) Evalua-
tion of chromatic cues for trapping Bactrocera tau. Pest Man-
agement Science 73: 217–222.
Liang DH, Chen R & Zhou Q (2016) Responsiveness of Bactro-
cera dorsalis (Hendel) adult to different colours and odorants.
Life Science Research 20: 36–39.
Lin YY, Tao J, Zeng L & Lu YY (2012) Cuticular penetration of b-
cypermethrin in insecticide-susceptible and resistant strains of
Bactrocera dorsalis. Pesticide Biochemistry and Physiology 103:
189–193.
Liu JH & Ye H (2006) Effects of light, temperature and humidity
on the flight activities of the oriental fruit fly, Bactrocera dor-
salis. Chinese Bulletin of Entomology 43: 211–214.
Mainali BP & Lim UT (2010) Circular yellow sticky trap with
black background enhances attraction of Frankliniella occiden-
talis (Pergande) (Thysanoptera: Thripidae). Applied Entomo-
logy and Zoology 45: 207–213.
Meyer HW (1978) Phototaxis in the walking male and female fly
(Calliphora erythrocephala Meig.). Journal of Comparative
Physiology 123: 307–314.
Meyer-Rochow VB & Lau TF (2008) Sexual dimorphism in the
compound eye of the moth Operophtera brumata (Lepi-
doptera, Geometridae). Invertebrate Biology 127: 201–216.
112 Liu et al.
Miller EC, Swanson AB, Phillips DH, Fletcher TL, Liem A &
Miller JA (1983) Structure-activity studies of the carcinogenici-
ties in the mouse and rat of some naturally occurring and syn-
thetic alkenyl benzene derivatives related to safrole and
estragole. Cancer Research 43: 1124–1134.
Okada Y, Tomioka K & Chiba Y (1991) Circadian phase-response
curves for light in nymphal and adult crickets, Gryllus bimacu-
latus. Journal of Insect Physiology 37: 583–590.
Pagadala DKJ, Christine MW, John C, Michael AB & Toby
JAB (2012) Isolation and identification of host cues from
Mango, Mangifera indica, that attract gravid female oriental
fruit fly, Bactrocera dorsalis. Journal of Chemical Ecology 38:
361–369.
Prokopy RJ & Owens ED (1983) Visual detection of plants by her-
bivorous insects. Annual Review of Entomology 28: 337–364.
Shelly TE (1997) Selection for non-responsiveness to methyl
eugenol in male oriental fruit flies (Diptera: Tephritidae). Flor-
ida Entomologist 80: 248–253.
Shen GM, Wang XN, Dou W & Wang JJ (2012) Biochemical and
molecular characterization of acetylcholinesterase in four field
populations of Bactrocera dorsalis (Hendel) (Diptera: Tephriti-
dae). Pest Management Science 68: 1553–1563.
Shimoda M & Honda KI (2013) Insect reactions to light and its
application to pest management. Applied Entomology and
Zoology 48: 413–421.
Silva FS, Silva AA & Reblo JMM (2016) An evaluation of light-
emitting diode (LED) traps at capturing phlebotomine sand
flies (Diptera: Psychodidae) in a livestock area in Brazil. Jour-
nal of Medical Entomology 53: 634–638.
Smith RL, Adams TB, Doull J, Feron VJ, Goodman JI et al.
(2002) Safety assessment of allylalkoxybenzene derivatives used
as flavouring substances – methyl eugenol and estragole. Food
and Chemical Toxicology 40: 851–870.
Stalleicken J, Labhart T & Mouritsen H (2006) Physiological
characterization of the compound eye in monarch butterflies
with focus on the dorsal rim area. Journal of Comparative
Physiology A 192: 321–331.
Sun YX, Tian A, Zhang XB, Zhao ZG, Zhang ZW & Ma RY
(2014) Phototaxis of Grapholitha molesta (Lepidoptera:
Olethreutidae) to different light sources. Journal of Economic
Entomology 5: 1792–1799.
Tang LD, Han Y, Wu JH, Li P, Fu BL et al. (2015) Preference of
Megalurothrips usitatus (Thysanoptera: Thripidae) to different
colors and light-waves in lab. Plant Protection 41: 169–172.
Tariq K, Noor M, Saeed S & Zhang H (2015) The effect of ultravi-
olet-A radiation exposure on the reproductive ability, longev-
ity, and development of the Dialeurodes citri (Homoptera:
Aleyrodidae) F1 generation. Environmental Entomology 44:
1614–1618.
Tariq K, Noor M, Hori M, Ali A, Hussain A et al. (2017) Blue
light-induced immunosuppression in Bactrocera doraslis
adults, as a carryover effect of larval exposure. Bulletin of Ento-
mological Research 107: 734–741.
Theobald JC, Coates MM, Wcislo WT & Warrant EJ (2007)
Flight performance in night-flying sweat bees suffers at low
light levels. Journal of Experimental Biology 210: 4034–
4042.
Vargas RI & Prokopy R (2006) Attraction and feeding responses
of melon flies and oriental fruit flies (Diptera: Tephritidae) to
various protein baits with and without toxicants. Hawaiian
Entomological Society 38: 49–60.
Vargas RI, Stark JD & Prokopy RJ (1991) Response of oriental
fruit fly (Diptera: Tephritidae) and associated parasitoids
(Hymenoptera: Braconidae) to different-color spheres. Journal
of Economic Entomology 84: 1503–1507.
Venter GJ, Hermanides KG, Boikanyo SNB, Majatladi DM &
Morey L (2009) The effect of light trap height on the numbers
of Culicoides midges collected under field conditions in South
Africa. Veterinary Parasitology 166: 343–345.
Vontas J, Hernandez-Crespo P, Margaritopoulos JT, Ortego
F, Feng H et al. (2011) Insecticide resistance in tephritid
flies. Pesticide Biochemistry and Physiology 100: 199–205.
Walcott B (1969) Movement of retinula cells in insect eyes on
light adaptation. Nature 223: 971–972.
Warrant E, Kelber A, Gislen A, Greiner B, Ribi W & Wcislo W
(2004) Nocturnal vision and landmark orientation in a tropical
halictid bee. Current Biology 14: 1309–1318.
Wei G, Zhang Q, Zhou M & Wu M (2000) Studies on the photo-
taxis of Helicoverpa armigera (H€ ubner). Acta Biophysica Sinica
16: 89–95.
Wu WY, Chen YP & Yang EC (2007) Chromatic cues to trap the
oriental fruit fly, Bactrocera dorsalis. Journal of Insect Physio-
logy 53: 509–516.
Xue HW & Wu WJ (2013) Preferences of Bactrocera cucurbitae
(Diptera: Tephritidae) to different colors: a quantitative inves-
tigation using virtual wavelength. Acta Entomologica Sinica
56: 161–166.
Yang ZX, Sha YC, Yuan LC, Zhang D, Yang SL et al. (2011)
Attraction effect of different colors and fruits for Bactrocera
dorsalis adults. Journal of Southwest University 33: 64–66.
Yoon JB, Nomura M & Ishikura S (2012) Analysis of the flight
activity of the cotton bollworm Helicoverpa armigera (H€ ubner)
(Lepidoptera: Noctuidae) under yellow LED lighting. Japanese
Journal of Applied Entomology and Zoology 56: 103–110.
Yu HL (2011) The Phototaxis of Grapholita molesta Busck and
the Effect of Green Light on Its Biological Characteristics. Mas-
ter’s Thesis, Northwest Agriculture and Forestry University,
Yangling, Shaanxi, China.
Zheng WW, Zhu CP, Peng T & Zhang HY (2012) Odorant recep-
tor co-receptor Orco is upregulated by methyl eugenol in male
Bactrocera dorsalis (Diptera: Tephritidae). Journal of Insect
Physiology 58: 1122–1127.
Zhou CQ, Chen HD & Lin PQ (1995) Comparison of the impact
of temperature, humidity and photoperiod on the population
reproductivity of three fruit flies. Acta Scientiarum Naturalium
University Sunyatseni 34: 68–75.
Zhu JL, Zhu W, Liu HA, Lu UR & Zhang CM (2014) Effects of
LED lights on phototaxis and reproduction of small brown
planthopper. Jiangsu Journal of Agriculture Science 30: 508–
513.