ARTICLE IN PRESS

Journal of Insect Physiology 53 (2007) 509–516

www.elsevier.com/locate/jinsphys

Chromatic cues to trap the oriental fruit fly, Bactrocera dorsalis

Wen-Yen Wua, Yu-Po Chena, En-Cheng Yanga,b,
a
Department of Entomology, National Chung Hsing University, Taichung, Taiwan
b
Laboratory of Insect Neurobiology, Department of Entomology, National Taiwan University, Taipei 10617, Taiwan
Received 27 December 2005; received in revised form 5 February 2007; accepted 5 February 2007

Abstract

Various colors have been used as visual cues to trap insect pests. For example, yellow traps for monitoring and control of the oriental
fruit fly (Bactrocera dorsalis) have been in use for a very long time. However, the chromatic cue of using color traps has never been
meticulously investigated. In this study, the spectral sensitivities of the photoreceptors in the compound eyes of B. dorsalis were measured
intracellularly, and the theory of receptor quantum catch was applied to study the chromatic cue of fly attracting. Responses to five
wavelength categories with peak wavelengths of 370, 380, 490, and 510 nm, and one with dual peaks at 350 and 490 nm were recorded.
Based on spectral sensitivities, six colored papers were chosen to test the color preference of the fly, and an additional UV preference test
was done to confirm the effect of the UV stimuli. It was concluded that UV and green stimuli (spectra: 300–380 nm and 500–570 nm)
would enhance the attractiveness of a colored paper to the oriental fruit fly, and blue stimuli (380–500 nm) would diminish the
attractiveness.
r 2007 Elsevier Ltd. All rights reserved.

Keywords: Spectral sensitivity; Color preference; Oriental fruit fly; Bactrocera dorsalis

1. Introduction Liu, 1981), and numerous lure-and-kill traps with yellow

Several species of fruit flies (Tephritidae) are major and
widespread destructive pests in agriculture around the
world (Christenson and Foote, 1960). Their oviposition
causes serious damage to fruit crops, costing millions of
dollars every year. To control and monitor these pests,
many studies have been devoted to trap-designs (Kat-
soyannos, 1989; Roessler, 1989; Heath et al., 1993). Since
host-finding is the indispensable stage for discovering a
suitable place for oviposition, and since it is strongly
influenced by both visual and olfactory cues (Prokopy and
Owens, 1983; Prokey et al., 1990), the best strategy to
attract these flies is by using these cues to develop effective
traps to lure them.
The oriental fruit fly, Bactrocera dorsalis, is the main
fruit fly pest in the Pacific Rim (Haramoto and Bess, 1970;
Corresponding author. Laboratory of Insect Neurobiology, Depart-
ment of Entomology, National Taiwan University, Taipei 10617, Taiwan.
Tel.: +886 2 33669640; fax: +886 2 33652092.
E-mail address: ecyang@ntu.edu.tw (E.-C. Yang).
colored surfaces have been developed and applied in the
field, such as yellow sticky papers and the famous male-
specific methyl eugenol-baited traps (Cornelius et al., 1999;
Alyokhin et al., 2000; Chen and Dong, 2001). Both Vargas
et al. (1991) and Cornelius et al. (1999) demonstrated that
yellow fruit-mimicking spheres were an excellent device to
lure the oriental fruit fly, and Alyokhin et al. (2000)
demonstrated that the visual cues of a yellow fruit-
mimicking sphere trap could increase the attractiveness of
hydrolyzed liquid protein odor. It has been shown that
visual and olfactory cues can function separately and have
a complementary effect to each other for attracting flies
(Jang and Light, 1991; Vargas et al., 1991; Cornelius et al.,
1999; Alyokhin et al., 2000; Corneluis et al., 2000a, b).
Thus, to optimize lure-and-kill traps, these cues must be
investigated carefully using experiments based on neu-
roethological aspects. However, most of these studies were
done only on olfactory cues for tephritids (Light et al.,
1992; Raptopoulos et al., 1995). Also, it should be noted
that using a color name as a reference to describe a fly’s
preference is not correct, since our color vision is quite

0022-1910/$ - see front matter r 2007 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jinsphys.2007.02.003
 ARTICLE IN PRESS
510 W.-Y. Wu et al. / Journal of Insect Physiology 53 (2007) 509–516
different from that of an insect, and it is better to define
the fly’s chromatic preference using the insect visible spec-
trum. In this paper, we report the wavelength preference
of the oriental fruit fly based on the evidence from
electrophysiological recordings complete with behavioral
demonstration.
Since the host-finding behavior is closely related to mate-
finding and oviposition behaviors in fruit flies (Prokopy
and Owens, 1983), it can be assumed that the inherent
wavelength preference is also closely related to host-finding
behavior. From a physiological perspective, the chromatic
cues of host-finding behavior can be taken as a different
proportion of stimulation for each type of photoreceptor.
Thus, selecting the chromatic stimuli according to the
spectral sensitivity of the photoreceptors for testing the
fly’s wavelength preference will yield more specific results
than studies referring to the host fruit of the fly only
(Vargas et al., 1991; Cornelius et al., 1999). We tested the
fly’s preference for colored papers that were selected based
on the spectral sensitivities of a fly’s photoreceptors, and
analyzed the results to reveal possible chromatic cues.
2. Materials and methods
2.1. Animals
Experiments were carried out on both male and female
adult oriental fruit flies, B. dorsalis, derived from a
laboratory stock maintained at 2871 1C in a rearing room
with 12:12 (L:D) photoperiod and fed with the peptone–
sugar mixture.
2.2. Spectral sensitivity measurement
Before electrophysiological manipulation, the experi-
mental fly was immobilized by placing it in a freezer with
chipped ice at 4 1C for 15–30 min. Then the head of the fly
was mounted on a brass pedestal, with a beeswax/rosin
(3:1) mixture, so that the head and thorax were rigidly
secured, and the abdomen was free to perform ventilatory
movements. A small window was cut in the dorsal–lateral
part of the left compound eye to expose the retinula for
inserting the recording microelectrode, and a silver wire
was inserted into the thorax as the indifferent electrode.
The damaged surfaces were covered immediately with
vaseline to prevent drying. The brass pedestal with the
mounted animal was placed on a metal plate with the
animal’s head at the center of a Cardan arm perimeter,
which was mounted with an optical fiber-guided stimula-
ting light source.
The microelectrode was made of microfilament alumi-
nosilicate capillary glass (O.D. ¼ 1.0 mm, I.D. ¼ 0.68 mm,
AF100-68-10, Sutter Instrument Co.) pulled on a Flaming-
Brown microelectrode puller (P-97 Flaming/Brown Micro-
pipette Puller, Sutter Instrument Co.), and had a resistance
of 140–160 MO when filled with 1 M lithium chloride
solution. Using a micromanipulator (MWS-32, Narishige
Scientific Instrument Lab.), the microelectrode was ad-
justed and lowered vertically to insert into the retinula
through the small window in the compound eye. After
inserting the microelectrode, the fly was dark-adapted for
at least 10 min. The following manipulations were per-
formed in the dark room to keep the fly dark-adapted. The
microelectrode was advanced meticulously, and a gentle
tapping was performed when the tip of microelectrode was
going to penetrating through the cell membrane. Each time
the baseline potential shifted, a series of flash stimuli were
used to check if the tip of microelectrode was impaling the
photoreceptor. If a depolarization response was detected,
the Cardan arm perimeter would be adjusted to align with
the visual axis of the recorded cell by obtaining the
maximum response. The recorded cell would be further
verified as a photoreceptor that the cell had a depolarized
graded response waveform, which was composed of an
initial on-transient depolarizing peak and a sustained
plateau, to an on-axis saturated ‘‘white light’’ stimulus of
200 ms duration. The signals measured by the microelec-
trode were preamplified 10  by an amplifier (Neuropobe,
Model 1600, A-M Systems Inc.) and then sent via a 12-bit
multifunction data acquisition system (PCI-6024E,
National Instruments) to an IBM compatible PC.
A Xenon-short arc lamp (XBO 1000W/HS/OFR,
OSRAM) was used as the stimulating light source. The
polychromatic ‘‘white light’’ was guided to pass a quartz
circular variable neutral-density wedge filter (Acton Re-
search Co.), which was capable of varying the intensity of
the light over a range of approximately 3 log units, and
then sent into the monochromator (SP-150-M with 150-
030-300 grating, Acton Research Co.). The separated
monochromatic light with half-band widths below 10 nm
was controlled by a magnetic shutter (SH-150, Acton
Research Co.) to form the flash stimulation with 20 ms
duration and was then guided by a UV-VIS fiber optic
bundle (LG-455-020-3, Acton Research Co.) to project on
the recorded eye. Since the terminal of the fiber optic
bundle with a diameter of approximately 1.5 mm was
230 mm away from the recorded eye, the terminal of the
fiber optic bundle therefore subtended an angle less than
0.371 at the eye.
The optic instruments and the multifunction data
acquisition system were controlled by a program developed
by LabVIEW software (ver. 6i, National Instruments) and
executed on the PC. Thus, it could produce equal quanta of
flux stimuli at 41 wavelengths from 300 to 700 nm, with
10 nm steps, and record the respective electrophysiological
responses from the photoreceptor. The amplitude of the
recorded responses were measured on-line and calculated
as the spectral sensitivity by the same program. All the
measured spectral sensitivity curves were calculated as the
mean of squared errors with the theoretical visual pigment
absorption curves, which were derived from the nomogram
by Ebrey and Honig (1977), at 41 peak wavelengths from
300 to 700 nm. The lmax was determined at the wavelength
with the minimum mean square error.
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W.-Y. Wu et al. / Journal of Insect Physiology 53 (2007) 509–516 511

2.3. Color preference test
To test the color preference of the oriental fruit fly, 6
colored papers (black, white, green, yellow, orange and
red) were chosen according to the spectral sensitivities of
the photoreceptors. Reflectance spectra of these colored
papers were obtained with a spectrometer (S2000, Ocean
Optics Inc.) compared with a reflectance standard (Spec-
tralon diffuse white standard, Labsphere, Inc.). The black
colored paper with almost no reflectance was chosen as the
test control. Yellow, orange and red colored papers were
chosen to test the effect of different stimuli on the
sensitivity peaks between 500 and 560 nm. The green and
white colored papers were chosen to test the effect of strong
and no UV stimulus (Fig. 1).
120
100
80
Reflectance (%)
Experiments were conducted on sunny days from April
to June 2004 on National Chung Hsing University campus.
Tests were performed in a dodecagon maze (Fig. 2)
outdoors, with two sets of colored papers attached
symmetrically on the side walls in random orders and with
each colored paper separated with opaque black plastic
plating. To allow the flies to see all 6 colored papers in any
horizontal direction, the tested flies restricted in a
transparent transport container were put at the center of
the maze before they were set free. After 3–5 min, the
container was opened to set the files free and allowed them
to move and fly freely to lower any possible stress effect.
The number of flies in each inside-chamber was counted
after 30 min. During the tests, the dodecagon maze with a
transparent top was placed in the shade of trees in order
to prevent any possible effects of polarization from
incident light.
Since it has been reported that the ERG of the fly may be
subject to age-related changes (Loew, 1975), and that
visual attractive cues for tephritid flies may differ between
sexes (Sivinski, 1990), the flies were tested separately for
different ages and sex with three replicates. Both male and
female flies from 1 to 20 days old from eclosion were tested.

Each test was performed on at least 30 flies; no fly was

60 tested more than once.

40 2.4. UV preference test

20
According to the result of color preference test (see
below and Fig. 8), the white colored paper with a high UV
reflectance was less attractive to the oriental fruit fly than
0
the green colored paper and even less attractive than the
300 400 500 600 700
black colored paper, a further test was performed to
Wavelength (nm)
evaluate the affect of UV stimuli on the color preference.
Fig. 1. Reflectance spectra of colored papers used in the color preference Twelve white colored papers were attached onto the inside
test. walls of the dodecagon maze as the above description. On

Fig. 2. Sketch of the dodecagon maze with 12 inside-chambers used for simultaneously comparing the attractiveness of different colored papers to oriental
fruit flies. The numbers indicate the dimensions of the maze and inside-chambers in centimeters. The structures of the inside-chambers are identically, and
the right illustration indicates the dimension of one of the inside-chambers marked in dark-gray color.
 ARTICLE IN PRESS
512 W.-Y. Wu et al. / Journal of Insect Physiology 53 (2007) 509–516

the top of the six chambers of the maze, UV-cut

transparent plastic plates servicing as UV cut-off filters 100
were applied to produce white colored paper without UV

Relative Sensitivity (%)

reflection below 380 nm, but the plates still had more than
80% transmission within 390–700 nm. The other six 75
chambers without the transparent plastic plates were
illuminated by normal light to the white colored papers,
50
and thus providing higher UV reflection from the papers.
All the six UV-cut chambers were separated by the normal
illuminated chambers. Therefore, the setup could provide
25
dual-choice of white colored papers with and without UV
illumination below 380 nm. The experimental procedure
followed the color preference test. Because the color
0
preference was not significantly different between the sexes 300 400 500 600 700
and the different ages (see color preference test result), the Wavelength (nm)
tests were performed with more than 25 males and 25
females with the age of 7 days old together at the same Fig. 3. The relative spectral sensitivity curve of the oriental fruit fly
time. Each test was performed with at least 50 flies; flies 370 nm photoreceptor (data averaged from three measurements of cell 15,
the scatter bar gives the standard error). The light gray dash line shows the
were tested only once. theoretical visual pigment absorption curve with the peak wavelength at
370 nm. The gray dot line shows the spectral sensitivity curve of the R7p
3. Results photoreceptor in the dipteran retinula (Hardie, 1986).

3.1. Spectral sensitivities of the compound eye

photoreceptors of the oriental fruit fly 100

Responses to five wavelength categories were measured

Relative Sensitivity (%)

from 37 cells in the dorsal–frontal part of the left 75

compound eye of the oriental fruit fly in dark adaptation.
The maximum intracellular response of the oriental fruit fly
photoreceptors was generally up to 70 mV. In some
recordings, however, such as in the 380 nm and the
510 nm photoreceptors, the recorded maximum response
could only reach about 25–40 mV. All recordings were
validated as intracellular by means of characteristics such
as the baseline drop during penetration of the cell
membrane (35 to 60 mV) and the transient-tonic
depolarized graded response waveform to a sufficient light
stimulus. The low maximum responses of these recordings
could have been caused by the filtering effects of the special
histological superposition condition (Wu et al., 1985), or
more likely the result of a microelectrode location distant
from the cell body of the photoreceptor. According to their
lmax of the spectral sensitivity curve, these photoreceptors
were grouped and termed as 370, 380, 490, 510 nm and
dual-peak photoreceptors. Further details of these photo-
receptors are described as follows:
370 nm photoreceptor: A single cell, with maximum
spectral sensitivity at 370 nm, was measured with three
measurements, one bi-directional run (300–700 nm and
700–300 nm) and another uni-directional run (from 300 to
700 nm). The black solid line of Fig. 3 shows that the
sensitive range of this cell was mainly located in the UV
range of 300–400 nm. The fitting procedure also demon-
strated that the curve was best fitted by the theoretical
visual pigment absorption curve of 370 nm visual pigment.
380 nm photoreceptor: The spectral sensitivity curve of
the 380 nm photoreceptor was similar to that of the 370 nm
50
25
0
300 400 500 600 700
Wavelength (nm)
Fig. 4. The relative spectral sensitivity curve of the oriental fruit fly
380 nm photoreceptor (data averaged from two measurements of cell 23).
The light gray dash line shows the theoretical visual pigment absorption
curve with the peak wavelength at 380 nm. The gray dot line shows the
spectral sensitivity curve of the R7y photoreceptor in the dipteran retinula
(Hardie, 1986).
photoreceptor in the UV range (Fig. 4), and formed a
plateau shoulder at 440–520 nm with sensitivity of
about 40% of the peak. The peak of the sensitivity curve
was at 370 nm, but due to the fact that the averaged curve
is best fitted with the theoretical visual pigment absorption
curve of 380 nm visual pigment (Fig. 4), this type is
therefore referred to as the 380 nm photoreceptor. The
cell was measured successfully with two measurements
(bi-directional run).
490 nm photoreceptor: Three measurements were success-
fully made on the 490 nm photoreceptor (one bi-directional
and another uni-directional run). This photoreceptor
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W.-Y. Wu et al. / Journal of Insect Physiology 53 (2007) 509–516
possesses a simple spectral sensitivity peak at 490 nm
(Fig. 5), and the spectral sensitivity curve is best fitted with
the theoretical visual pigment absorption curve of 490 nm
visual pigment.
510 nm photoreceptor: The shape of the spectral sensitiv-
ity curve of the 510 nm photoreceptor appeared to be like a
‘‘screened’’ curve of the 490 nm photoreceptor (Fig. 6).
Therefore, the spectral sensitivity peak was ‘‘shifted’’ to the
range of 480–550 nm, and the curve was best fitted with the
theoretical visual pigment absorption curve of 510 nm
visual pigment (Fig. 6). The cell was also measured with
100
Relative Sensitivity (%)
75
50
25
0 Po0.0001; ANOVA Procedure of SASs). Therefore the
300 400 500 600
Wavelength (nm)
Fig. 5. The relative spectral sensitivity curve of the oriental fruit fly
490 nm photoreceptor (data averaged from three measurements of cell 40,
the scatter bar gives the standard error). The light gray dash line shows the
theoretical visual pigment absorption curve with the peak wavelength at
490 nm. The gray dot line shows the spectral sensitivity curve of the R8p
photoreceptor in the dipteran retinula (Hardie, 1986).
100
Relative Sensitivity (%)
75
50
25
0 0
300 400 500 600
Wavelength (nm)
Fig. 6. The relative spectral sensitivity curve of the oriental fruit fly
510 nm photoreceptor (data averaged from six measurements of cell 14,
the scatter bar gives the standard error). The light gray dash line shows the
theoretical visual pigment absorption curve with the peak wavelength at
510 nm. The gray dot line shows the spectral sensitivity curve of the R8y
photoreceptor in the dipteran retinula (Hardie, 1986).
513
three repeats (one bi-directional and another uni-direc-
tional run).
Dual-peak photoreceptor: These photoreceptors were the
photoreceptor most often encountered in 33 cells in the
retinula. The spectral sensitivity curves of these cells were
composed of two major peaks at the UV (330–370 nm) and
the blue-green (430–510 nm) range. The band widths of the
peaks and the amplitude ratios between the peaks showed
some varieties. Fig. 7 shows the spectral sensitivity curve
averaged from the measurement of 7 cells, the lmax was
obtained at 350 and 490 nm by fitting with the theoretical
visual pigment absorption curve.
3.2. Color preference of the oriental fruit fly
The preferred choice rates were obtained from three
replicates of each age and sex. These rates were not
significantly related to either age or sex when analyzed with
a three-way ANOVA test, but they were significantly
related to the color of the colored papers (F ¼ 88.77;
df ¼ 5; Po0.0001; ANOVA Procedure of SASs). Choice
rates were also not significantly related to the mature status
of the flies (1–7 days old flies and older flies) when analyzed
with a two-way ANOVA test, but were significantly related
to the color of the colored papers (F ¼ 73.88; df ¼ 5;
700 data from both sexes and all ages were combined for
further analyses. Fig. 8 shows the mean choice rates for the
colored papers from 120 replicates (three replicates for 20
ages and two sexes). The green, yellow and orange colored
papers, with an oriental fruit fly attracting rate of
(Mean7SE) 0.2265270.00766, 0.2194170.00826, and
0.1913270.00704, respectively, were significantly more
attractive than the red, black, or white colored papers
100
Relative Sensitivity (%)
75
50
25
700 300 400 500 600 700
Wavelength (nm)
Fig. 7. The relative spectral sensitivity curve of the oriental fruit fly dual-
peak photoreceptors (data averaged from seven cells, the scatter bar gives
the standard error). The light gray dash lines show the theoretical visual
pigment absorption curves with the peak wavelength at 350 and 490 nm.
The gray dot line shows the spectral sensitivity curve of the R1–6
photoreceptors in the dipteran retinula (Hardie, 1986).

514 W.-Y. Wu et al. / Journal of Insect Physiology 53 (2007) 509–516
0.25 a 370 and 380 nm photoreceptors: The spectral sensitivity
a,b
b the R7p photoreceptor obtained from other flies (Fig. 2;
0.20
c
Attraction Rate
0.15
0.10
0.05
0.00
Yellow Green Orange Red
Color Paper
Fig. 8. Attraction (Mean+SE) of different colored papers to the oriental
fruit fly. The same lowercase letters above the bars in (a–c) indicate no
significant difference (F ¼ 73.61; df ¼ 5; Po0.0001; ANOVA Procedure
and Tukey’s Studentized Range (HSD) Test of SASs).
(Fig. 8). The white colored paper with the 0.079970.00504
oriental fruit fly attracting rate was significantly less
attractive to the oriental fruit fly than the red and black
colored paper with an attracting rate of 0.1487470.00707
and 0.1341170.00578, respectively (Fig. 8; F ¼ 73.61;
df ¼ 5; Po0.0001; ANOVA Procedure of SASs).
3.3. UV preference test
The choice rates were obtained from 3 replicates, which
showed that the white colored papers with the UV
illumination below 380 nm were significantly more attrac-
tive than the white colored papers without the UV
illumination (Mean7SE: 0.4670.03055 vs. 0.246677
0.04807; F ¼ 13.17; df ¼ 1; Po0.05; ANOVA Procedure
and Tukey’s Studentized Range (HSD) Test of SASs).
4. Discussion
4.1. Spectral sensitivity of the photoreceptors
Dual-peak photoreceptor: The dual-peak photoreceptors
have a similar action spectrum to the R1–6 photoreceptors
found in other dipteran flies (Fig. 7; Horridge and Mimura,
1975; Horridge et al., 1975; Hardie, 1979, 1986; Hardie et
al., 1979). The antenna pigment hypothesis proposed by
Kirschfeld et al. (1977) suggested that the blue-green peak
was from the intrinsic blue-green sensitive rhodopsins and
that the UV peak was due to the UV sensitizing pigments
transferring their absorbed energy to the rhodopsins. This
would then make the action spectra of these photorecep-
tors become broadly distributed with two peak wave-
lengths. Variation of the spectral sensitivity curves can be
assumed to be the result of self-absorption or electric
coupling effects (Manzel, 1975).
ARTICLE IN PRESS
curve of the 370 nm photoreceptor is similar to the curve of
Horridge et al., 1975; Hardie, 1979, 1986; Hardie et al.,
1979). It is very likely that the sensitivity range of the
c 370 nm photoreceptor may be elicited by a single visual
pigment with its lmax at 370 nm. In contrast, the 380 nm
photoreceptor has a more complicated spectral sensitivity
d curve, in which a shoulder was observed from 440 to
520 nm. This curve is also similar to the R7y photoreceptor
of other dipteran flies (Fig. 3; Horridge et al., 1975; Hardie
et al., 1979; Hardie, 1986). According to Hardie (1986), the
mechanism that elicits these action spectra of the photo-
receptors may possibly be caused by the fact that the blue
Black White sensitive rhodopsins (e.g. xanthopsin, lmax ¼ 430 nm) of
the cell were greatly hindered by the blue screening pigment
(e.g. carotenoid). However, the cells possess a UV
sensitizing pigment similar to the dual-peak photoreceptors
and the absorbed energy of the unscreened sensitizing
pigments can transfer to the blue sensitive rhodopsins and
cause the UV peak with the blue shoulder in the action
spectra of these cells. Therefore, the 380 nm photoreceptor
has a complicated action spectrum, and the summation of
UV sensitizing pigment and the ‘‘hindered’’ blue sensitive
rhodopsins makes the spectral sensitivity curve best fit with
the 380 nm theoretical visual pigment absorption curve. In
addition, the block effects of the blue screening pigments in
the 380 nm photoreceptor of the oriental fruit fly would be
less than those in the R7y photoreceptor of other dipteran
flies, as the 380 nm photoreceptor was more sensitive than
the R7y photoreceptor within the range between 420 and
510 nm.
490 and 510 nm photoreceptors: The spectral sensitivity
curves of 490 and 510 nm photoreceptors are similar to the
R8p and R8y photoreceptors of other dipteran flies (Figs. 4
and 5; Hardie et al., 1979; Hardie, 1986). It is possible that
the 490 and 510 nm photoreceptors of oriental fruit fly are
the R8 cells described by Wu et al. (1985). Because the
rhabdomere of the R8 cell is beneath the rhabdomere of the
R7 cell in the oriental fruit fly (Wu et al., 1985), the 370 and
380 nm photoreceptors may be considered as the R7 cells. If
this is the case, the spectral reception of the 490 and 510 nm
photoreceptors should be filtered by the 370 and 380 nm
photoreceptors, respectively. Because the spectral absorp-
tions of 490 and 370 nm photoreceptors are not over-
lapping in their dominant peaks, the 490 nm photoreceptor
still keeps its spectral sensitivity in the blue-green range of
spectrum. On the other hand, because the spectral
absorption of the 380 nm photoreceptor and the spectral
sensitivity of the 510 nm photoreceptor overlapped, light
absorption by the UV sensitizing pigment and the blue-
green sensitive rhodopsin of the 510 nm photoreceptor is
hindered by the 380 nm photoreceptor. Consequently, UV
sensitivity is diminished as a small sensitive peak, and the
blue-green sensitivity is also altered, so that the spectral
sensitivity curve fits best with the 510 nm theoretical visual
pigment absorption curve.
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W.-Y. Wu et al. / Journal of Insect Physiology 53 (2007) 509–516
4.2. Color preference of the oriental fruit fly
Previous studies on another tephritid pest, the apple
maggot fly (Rhagoletis pomonella), demonstrated that the
intensity contrast of dark host against bright background
plays a more important role in the fly’s host detection than
host colour (Agee, 1985; Owens and Prokopy, 1986).
Contrarily, a bright host (e.g. colored with yellow or white)
is more attractive to the oriental fruit fly when tested with
different-color spheres (Vargas et al., 1991), suggesting that
different strategies are used for seeking host targets
between the two fruit fly species.
When comparing the reflectance spectra of the colored
papers and their attractiveness for the oriental fruit fly
(Figs. 1 and 8), it is evident that several spectral cues are
involved with the attractiveness. The green colored paper
that showed the higher attractiveness among the colored
papers has a broader and higher reflectance, indicating that
chromatic stimulation of the spectra within 500–570 nm
contains crucial cues for attracting the oriental fruit fly.
This is also supported by the fact that orange colored paper
with a reflectance spectrum slightly higher than that of red
colored paper, in the range of 470–630 nm, has a
significantly stronger attractiveness to the flies than the
red colored paper. In addition, the white colored paper
used in our experiment has the broadest reflectance
spectrum but the lowest attractiveness to the flies, even
less than the black colored paper, and the green colored
paper has a similar attractive effect as the yellow colored
paper, which has a much higher reflectance within
500–570 nm than the green colored paper. Thus, it is
reasonable to hypothesize that there is a negative cue in the
spectrum to reduce the attractiveness.
The UV-sensitive peaks seen in the recorded spectral
sensitivity curves of the fly initially suggested that negative
cues might be present in this spectral region. However, the
preference test indicated that UV reflection enhanced
attractiveness, as did the ‘‘green’’ reflection (500–570 nm).
The UV-positive effect could be attributed to the UV
sensitivity of the dual-peak photoreceptors as well as the
370 nm and the 380 nm photoreceptors. A final possibility
for the location of the negative cue could be within the
visual sensitive range of 380–500 nm, for which no
corresponding photoreceptor was recorded in this study.
This negative effect could be caused by opponent mechan-
isms, or related to an undiscovered blue photoreceptor
(sensitivity range between 400 and 500 nm), as suggested by
a previous ERG study on the oriental fruit fly (Wu, 1989)
and the evidence shown in other dipteran flies (Hardie,
1986). The green-positive effect could be attributed to the
green sensitivity of the 510 nm photoreceptor, or the
related opponent mechanisms as mentioned above. Since
the action spectra of the photoreceptors overlapped each
other, genetic mutants such as photoreceptor-defective
strains or RNAi knock down of individual photopigments
would be required to confirm which photoreceptors were
actually involved with color preference. Nevertheless, the
515
chromatic cues of the color preference obtained by
applying the fundamental physiological information are
helpful in studying the visual ecology of the oriental fruit
fly, and provide a useful tool for developing pest control
methods. While various yellow traps have been widely used
in field trapping, most of the ‘‘yellow colors’’ were
determined by human perception without precisely quanti-
fying the action spectrum of the pest and concerning the
negative effect. Our results suggest that a trap with
sufficient reflection within 300–380 nm and 500–570 nm
but without reflection within 380–500 nm will be more
effective in attracting the oriental fruit fly.
Acknowledgment
We thank the anonymous reviewers for insightful
corrections and improvements. We are also most grateful
to Dr. Kuang-Hui Lu and his lab for their generous fly
stocks. This work was supported by a grant (94 AS-13.2.1-
BQ-BG) from the BAPHIQ and a grant (NSC 91-2313-B-
005-117) from National Science Council, Taiwan.
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