Journal of Integrative Agriculture
Advance Online Publication 2014
DOI：10.1016/S2095-3119(14)60863-7
Spectral Sensitivity of the Compound Eyes of Anomala corpulenta Motschulsky
(Coleoptera: Scarabaeoidea)1
JIANG Yue-li12, GUO Yu-yuan13, WU Yu-qing2, LI Tong2, DUAN Yun2, MIAO Jin2, GONG Zhong-jun2 and
HUANG Zhi-Juan2
1
College of Plant Protection, Northwest A&F University, Yangling, Shaanxi , 712100, P.R.China
2
Institute of Plant Protection, Henan Academy of Agricultural Sciences, Key Laboratory of Crop Pest Control of Henan Province, Zhengzhou 450002,
P.R China
3
State Key Laboratory for the Biology of Plant Diseases and Insect Pests, Institute of Plant Protection, Chinese Academy of Agricultural Sciences,
Beijing 100193, P.R.China
Abstract
The scarab beetle, Anomala corpulenta Motschulsky (Coleoptera: Scarabaeoidea, Rutelinae), is a widespread and destructive pest in
China. Vision is one of the most important means of acquiring information about the external environment. In order to contribute to the
understanding of the perception of visual stimuli in this species, the light sensitivity and spectral responses of the scarab beetle, A.
corpulenta, were measured by using an electroretinogram (ERG) technique. In total, 14 monochromatic light intensities, between 340
nm and 605 nm, were applied to the compound eyes of A. corpulenta under varying levels of adaptation to dark and light conditions.
The results showed that all light stimuli induced an ERG reaction, with varied amplitudes. The spectral sensitivity curve of
dark-adapted eyes showed one major peak (~400 nm; near- ultraviolet), a secondary peak (from 498 nm to 562 nm; yellow-green) and
a third peak at 460 nm. By contrast, in light-adapted eyes, only a near-UV peak was observed. From these results, we conclude that the
compound eye of A. corpulenta is likely to have at least three spectral types of photoreceptor. Significance of differences were also
recorded in the responses of male and female compound eyes, as well as diurnally and nocturnally. The amplitude of ERG in response
to white-light stimuli varied with the light intensity: the stronger the luminance, the higher the ERG value. This suggests that the
compound eye of A. corpulenta adapts quickly to changing light conditions, enabling A. corpulenta to maintain nocturnal activities.
Key words: A. corpulenta, electroretinogram, insect vision, spectral sensitivity, light intensity.
INTRODUCTION
For insects, vision is one of the most important means of acquiring information about the external environment: it
plays a significant role in finding hosts, intraspecific communication, foraging, escaping from predators and in
flight (Moericke 1955; Chapman et al. 1981; Dixon 1985; Klingauf 1987; Powell et al. 1995; Egelhaaf and Kern
2002). Finch and Collier (2000) showed that visual cues were central link of host location phytophagous insects,
1
JIANG Yue-li, Tel: +86 371 65738134, E-mail: [email protected]; Correspondence GUO Yu-yuan, Tel : +861062894786, E-mail: [email protected]
Correspondence WU Yu-qing, Tel: +86 371 65738134, E-mail: [email protected]
such as the butterfly Pieris rapae was found to react strongly to blue and blue-green light over long distances to
locate a suitable spawning host (Qin 1987), female fruit flies were more drawn to red ball-shaped traps the center of
which resembled yellow oranges (Cornelius et al. 1999). In addition, many phytophagous insects prefer yellow as a
cue to host location (Bernays and Chapman 1994). However, Frisch (1949) showed that the honeybee Apis
mellifera detects polarized natural light and uses it for spatial navigation. Therefore, it is important to get a more
comprehensive and clear understanding of the insect color vision.
The scarab beetle, A. corpulenta Motschulsky (Coleoptera: Scarabaeoidea, Rutelinae), is a widespread and
destructive pest in China (Wu 2001). The larvae feed on the underground portion of crops, with a preference for
peanuts and soybeans, whereas the adults feed on the leaves of various species of fruit and forest trees. The
exocuticle of this beetle has a brilliant metallic appearance, and selectively reflects left circularly polarized light, as
recorded for other jewel beetles (Sharma et al. 2009).
Insects are sensitive to various characteristics of light, such as its intensity, color and polarization (Wehner 1983;
Wernet et al. 2003; Horváth and Varjú2004). In addition, visual cues are important in intraspecific communication
and host location by beetles. Many studies on vision in the jewel beetles have focused on polarization (Horváth and
Varjú2004; Brady and Cummings 2010; Blahóet al. 2012). However, few have studied the presence of color
vision in, and impact of light intensity on, such beetles. Therefore, there is a need to further understand the
physiological basis of the color vision of jewel beetles, particularly the spectral sensitivity, and effects of light
intensity. Electrophysiological spectral sensitivity has been examined for many insect species, including the cabbage
root fly, Delia radicum (L.) (Brown and Anderson 1996), the cotton bollworm, Helicoverpa armigera (Hübner) (Wei et
al. 2002), the western flower thrip, Frankliniella occidentalis (Pergande) (Matteson et al. 1992), Homopteran species-the
glasshouse whitefly, Trialeurodes vaporariorum (Westwood) (Mellor et al. 1997), and the green peach aphid, Myzus
persicae (Sulz.) (Kirchner et al. 2005), the click beetles, Pyrophorus punctatissimus in Coleoptera (Lall et al. 2000), etc.
Most insects have two kinds of visual pigment, one that detects light at approximately 550 nm (yellow-green) and the
other that detects blue-violet UV light, at less than 480 nm. However, there are no data available regarding the
electrophysiological spectral sensitivity of the jewel beetles. In addition, mechanisms of adaptation to light intensity in A.
corpulenta are unclear.
Therefore, to investigate the spectral sensitivities and mechanisms of adaptation to light intensity of A.
corpulenta, the response of A. corpulenta to different light wavelengths and intensities, and the effects of gender
and circadian rhythm on them, were measured by means of electroretinogram (ERG) technique.
RESULTS
Form of the ERG
2
White light stimulation typically elicited the compound eye of A. corpulenta an ERG waveform that was a
monophasic negative change in potential (Fig. 1). As can be seen from the waveform, the lower interference signals
existed during stimulation. The ERG value was recorded as the negative ERG component. The greatest ERG
amplitude was 35 mV.
Spectral sensitivity
The spectra of ERG changes by the eyes of male and female A. corpulenta in response to light stimuli within the
monochromatic 340-605 nm wavelength were recorded under certain dark and light adaptation times. The results
showed that UV and most of the visible regions of the monochromatic light stimulus triggered an ERG reaction of
A. corpulenta compound eyes with light and dark conditions adaptation. The spectral sensitivity response curves
(Figs 2, 3 and 4) were obtained according to the size of the ERG amplitude.
The spectral sensitivity of the dark-adapted compound eye of both male and female A. corpulenta showed one
major distinct peak of sensitivity at 400 nm diurnally. The second distinct peak appeared at 524 nm (yellow-green).
No clear peak was found at other wavelengths (Fig. 2). When the eyes were tested nocturnally, the spectral
sensitivity curve showed three further peaks (Fig. 3). The main peak position was unchanged, whereas the
secondary peak position moved from 524 nm to 498 nm (yellow-green). The third peak occurred at 460 nm.
When the compound eyes were tested in the presence of adaptation to light, the spectral sensitivity curve of the
female eyes showed only one peak, at 400 nm (Fig. 4), whereas those of the males showed three peaks: a major
distinct peak at 400 nm, a second distinct peak at 498 nm (yellow-green) and a third peak at 460 nm. These peaks
were the same as, if smaller than, those recorded for the dark-adapted compound eye nocturnally.
By Using Mann-Whitney U test a comparison of mean spectral efficiency between the sexes failed to show any
significant differences on either day and night at corresponding wavelengths under light and dark adaptation
(P>0.05).
Circadian rhythm can also affect the spectral sensitivity of the compound eyes of A. corpulenta. As shown in
Figs 2 and 3, spectral sensitivity was stronger at night under dark-adaptation conditions than it was diurnally. There
were no significant differences in spectral efficiency between sexes (P>0.05), and therefore the data
from males
and females were combined. The Mann-Whitney U test revealed that the compound eye of A. corpulenta was
significantly more efficient in the peak of 400 nm (P<0.05), 460nm (P<0.05), 498nm (P<0.01), 538-582nm
(P<0.05) region at night under dark-adaptation conditions than it was diurnally. From 340-380 nm (P>0.05) ,
420-440 nm (P>0.05), 483nm (P>0.05), 524nm (P=0.05) and 605nm (P>0.05) the efficiencies of the day and nignt
were similar.
3
Changes in the light intensity reaction of the compound eye
By adjusting the neutral density filter, it was possible to stimulate the compound eyes with different intensities of
white light both diurnally and nocturnally. The light intensity reaction of compound eyes of male and female A.
corpulenta were recorded under dark-adaptation conditions (Figs. 5 and 6): the ERG value gradually increased with
increases in light intensity across a range of intensity (LogI=4.5~0). The increase was slow during initial
light-intensity levels, but increased more quickly once the light intensity reached a certain level. The increase was
not affected by gender or circadian rhythm, and the high-end platform of V- LogI reaction was not found yet
because of the limit of the light source.
DISCUSSION
According to the ERG, we found that the ERGs recorded from the compound eyes of A. corpulenta were similar to
that described in other insects, such as Curausius morosus ( Kugel 1977), Periplaneta americana (L.) (Walther
1958) and many species of Lepidoptera ( Eguchi et al. 1982).
The results of this study show that UV and most visible monochromatic stimuli can trigger an ERG reaction of
A. corpulenta compound eyes with light and dark conditions adaptation. Specially, we found spectral sensitivity of
A. corpulenta. The curves in Figs 2-4 suggest the presence of three peak regions of the spectral sensitivity system
of A. corpulenta: one in the near UV (400 nm), a broader green peak (498-562 nm) and the third peak at 460 nm.
With white adapting light, which reduces the sensitivity of any receptors in the visible range, the sensitivity curve
peaks at 400 nm. From these results, one can conclude that A. corpulenta has at least three types of photoreceptor.
Thus, this species is obviously similar in its visual system to other Coleoptera, such as North American fireflies
( Lall et al. 1980a; Lall et al. 1980b; Lall 1981)and Lepidoptera, such as Papilio protenor, Celastrina argiolus,
Minois dryas, Neope goschkevitschii, Celastrina argiolus, Ochlodes venata and Parnara guttata ( Eguchi et al.
1982) and Aglais urticae ( Steiner et al.1987).
Evidence of the near UV receptor (400 nm) of A. corpulenta comes from the results of both the dark adaptation
(both nocturnally and diurnally) and the light adaptation tests, because the peak at 400 nm was unchanged. During
the diurnal dark-adaptation test, the blue peak (at 460 nm) disappeared, but reappeared during the nocturnal
dark-adaptation test. This discrepancy in results might be because A. corpulenta are nocturnal insects and, therefore,
are most active during night and so have evolved a suitable dark-adapted visual system. With adaptation to white
light, which reduces the sensitivity of any receptors in the visible range, the clear sensitivity curve peaked only at
400 nm, indicating the existence of a UV receptor.
The near-UV spectral sensitivity recorded in A. corpulenta is a common feature of the compound eye of many
insects (Briscoe and Chittka 2001). It has been demonstrated that many insects use near-UV wavelengths in
4
navigation (Wehner 1976), and eliciting phototaxis of many insects mainly rely on the spectrum region, because it
is the most effective (Helen and Moray 1996). Some researchers suggested that the UV receptor might be optimally
suited to detec the open sky or to detect small objects (such as flying mates) against the bright sky(Menzel 1979). It
has been hypothesised that near-ultraviolet light serves to elicit escape reactions in the butterfly, Pieris (Scherer and
Kolb 1987).
For the green-sensitive visual pigment, a broader secondary peak in the spectral sensitivity curve, between
498 nm and 562 nm, was recorded from compound eyes of A. corpulenta under dark-adapted conditions. These
results are similar to those from other species of Coleoptera (Hasselmann 1962; Lall et al. 1982; Lin and Wu 1992).
Briscoe and Chittka (Briscoe and Chittka 2001) reported that green-sensitive photoreceptors are also a common
feature of the compound eyes of a variety of insects. Green-sensitive photoreceptors are quite effective region of
inducing some insects to green - yellow colors (Harris and Miller 1983; Prokopy et al.1983; Bernays and Chapman
1994). Phytophagous insects may use green-sensitive photoreceptors for host location (Bernays and Chapman
1994), and wide-field motion detection (Wehner 1981).
Vision is the main sense used for acquiring information from the external environment, enabling many species
of insect to find mates, obtain food, escape from predators and to fly (Egelhaaf and Kern 2002). In the current study,
we compared the visual spectral sensitivity and the color on the exoskeletal surface in both male and female A.
corpulenta. There was an approximate correlation between green color on the exoskeletal surface and the broad
green visual spectral sensitivity of this insect. However, the biological meaning of the phenomenon is hard to
explain. A similar phenomenon was reported in lycaenid butterflies (Michio and Kaoru 2008). The color vision of A.
corpulenta beetles might be more sensitive in discriminating conspecific signals from background light (contrast
enhancement) because this remarkable ability is known to mediate sexual signaling and mate choice in the scarab
beetle Chrysina gloriosa (Brady and Cummings 2010). However, additional work is required to investigate this
hypothesis further.
According to trichromatic theory of insect colour vision, and insects see three primary colours, green, blue and
UV (Briscoe and Chittka 2001). In the current study, A. corpulenta having 3 types of photoreceptors, it is
remarkable that the ERG of the click beetles (Coleoptera: Elateridae) was found to have just 2 spectral peaks in the
ERG (Lall et al. 2000), studies showed that some members of Coleoptera lost blue opsin (Jackowska 2007, Oba
2009). Blue photoreceptors are mainly reponseable for eliciting phototaxis of many insects, such as blue sticky
traps for monitoring Frankliniella bispinosa (Morgan) (Thysanoptera: Thripidae)(Childers et al. 1996). Further
studies are required, molecular evidence of 3 types of photosensitive receptors in A. corpulenta, and the function of
A. corpulenta photoreceptor.
A small sexes difference in spectral sensitivity curves was found in A. corpulenta beetles (Figs 2 and 3), but
there were no statistical differences between the sexes. A similar result was reported in Ephestia
cautella (Helen and Moray 1996), Trialeurodes vaporariorum and Encarsia formosa ( Mellor et al. 1997). It
5
showed that the sex differentiation was not obvious.
As other studies have shown, circadian rhythm can affect the spectral sensitivity of insects (Kral and Stelzl 1998;
Wei et al. 2002). In the current study, we found that circadian rhythm can affect the peak position of ERG. The
second distinct peak appeared from 524 nm during daylight hours to 498 nm during the night as a result of the
compound eyes becoming dark adapted. The peak at 460 nm appeared under dark-adapted conditions nocturnally
but not during daylight hours or under conditions of nocturnal light adaptation. The spectral sensitivity curves of
male and female A. corpulenta were higher nocturnally than diurnally, showing that the compound eyes of this
species are more sensitive during the night. Such a result supports the fact that these beetles are more active during
the night.
In addition, intensity response of A. corpulenta is also a significant founding. In certain intensity range of the
white-light, the stronger the luminance, the higher the ERG value of eyes from A. corpulenta. This suggests that the
compound eye of A. corpulenta has a strong ability to adapt to light, as also reported by Wei et al. (2002) and Yan
et al. (2007).
In the current study, the ERG technique was used to examine adaptive aspects of color vision in insects. This
technique is not capable of exploring the characteristics of receptors or pigments that play a role in color perception
and consequently it can not be used to investigate evolutionary aspects of the visual system. In order to improve our
understanding of the visual physiology and photosensitive mechanisms of A. corpulenta, further study on the
intracellular potential is necessary. Therefore, a combination of research at the molecular-, cellular- and
whole-eye-level approach, is required to further our understanding of the physiological mechanism of A. corpulenta
color vision.
CONCLUSION
In conclusion, the compound eye of A. corpulenta is sensitive to near-UV, green-yellow and blue light, circadian
rhythm can also affect the light sensitivity and spectral responses of the beetles. A. corpulenta is likely to have at
least three spectral types of photoreceptor. Its compound eyes have a strong ability to adapt to light. Specifically, in
part of the tested white-light intensity range, the stronger the luminance, the higher the ERG values recorded from
the compound eyes of this beetle.
MATERIALS AND METHODS
Insects
The A. corpulenta used in the study were taken from a natural population that had been collected during the
summer of 2012 in the Yuanyang Experimental Station (113.46°E; 35.08°N), Henan Academy of Agricultural
6
Sciences, Henan Province, China. The beetles were reared on tomato plants after collecting under the following
environmental conditions: 25℃, 70% relative humidity (RH) and a 16 h: 8 h light: dark photoperiod. The insects
used in the experiment were l-2-week-old adults.
Optical system
A 150-W high-pressure xenon lamp was used as a light source. The wavelength of stimulated light was obtained by
using an interference filter. The light passed first through a quartz heat-protecting glass, which absorbed the
infrared light, and then through a set of interference filters and neutral filters, which produced monochromatic light
with different intensities. The wavelengths of monochromatic light were 340, 360, 380, 400, 420, 440, 460, 483,
498, 524, 538, 562, 582 and 605 nm.
The optical path system comprised a quartz thermal-isolating glass, an interference filter, a neutral filter and
neutral wedge filter, which was used to adjust the light intensity. In addition, there was a quartz condenser lens,
which made parallel light sources converge on the focal plane of the light guide (inner diameter of 3 mm). The
shutter drive was controlled by a computer, which was used to trigger the shutter in real-time, to regulate the time
interval of light stimulating, and to enable the light beam to reach the compound eye through the light guide.
Intensity calibration
The neutral filter and neutral wedge filter were used to keep the light intensity of different monochromatic light the
same; this was monitored using a thermal photoelectric-coupled illuminance meter and a galvanometer [street
lighting photometer (Made by EVANS Electroselenium LTD, HALSTEAD ESSEX ENGLAND)]. Each duration of
light stimuli was set for 20 ms and the interval between stimuli was 40 s. In the light-intensity experiment, the light
intensity of white light was increased gradually logarithmically, with a grade of 4.5, 4, 3.5, 3, 2.5, 2, 1.5, 1, 0.5 and
0(Log I), by means of a number of neutral filters. The maximum intensity was at the Log 0. Each duration of light
stimuli was set for 20 ms and the interval between stimuli was 10 s.
Experimental procedures
Each insect was then mounted on a multidimensional adjustable ball-joint sample stage using low-melting-point
wax melted onto the abdomen-thorax. The head of the insect was carefully immobilized using melted wax. Under
an anatomical lens, a triangular microhole in the cornea of the left compound eye was made using microscissors;
Vaseline was used to seal the hole to prevent water loss before a recording electrode (see section 2.3.2) was
inserted into the left eye through the hole in the cornea.Try to keep a consistent depth when recording electrodes
were inserted into the eye every test. The reference electrode was placed in the thorax. Each glass microelectrode
was filled with 3 mol/L potassium acetate. The resistance of the said microelectrode was approximately 60 megohm.
7
The sample stage with the mounted insect was placed on the holder of the micro-electrode propeller. The signal,
which was amplified using a Preamplifier (Electrode amplifier L/M-1, UST-Medical-Electronic, Germany) was
monitored using an oscilloscope (PM3234 0-10，Philips) and the data saved to a computer. After insertion of the
electrode into the eye, the position in which the light elicited the maximum response of the eye was chosen for
stimuli application. The amplitude and duration of potential was used as a proxy for the size of the retinal cell
potential.
The spectral sensitivity was investigated under two different conditions: (i) dark adaptation, with an adaptation
time of 60 min; and (ii) light adaptation using white light, where the light intensity was approximately 100 lx and
the adaptation time was also 60 min. The effects of light intensity were investigated using dark-adapted conditions,
with an adaptation time of 60 min. Adaptation time refer to reported similar insect.(Lall et al. 2000)
Data analysis
Each ERG value from experimental recording in A. corpulenta was converted to a sensitivity value (Sn) using the
following Equation:
Sn=100×10-(logImax-logIn)%
where logImax is the equivalent strength of the maximum voltage response, logIn is the equivalent strength of
every voltage response ( Defrize et al. 2011).
Graphs were produced using the software origin 8.0. Mann-Whitney U tests (Grimm 1993) were applied to
determine differences in efficiency at corresponding wavelengths between sexes and between circadian rhythm of A.
corpulenta.
Acknowledgements
We especially thank Professor Guo-Shu Wei, Hebei Agricultural University, Baoding, China, for kindly providing
the equipment. We are also grateful to Doctor Xin-Cheng Zhao, Henan Agricultural University, Zhengzhou, China,
for assistance in modifying the article.The Special Fund for Agro-scientific Research in Public Interest (Project no.
201003025) and the Henan Academy of Agriculture Sciences Financial Budget (2012) funded this research.
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FIGURE LEGENDS
Fig. 1 A typical electroretinogram recording from A. corpulenta under white-light stimulus. A = negative sustained potential, B =
off-response.
Fig. 2 Spectral sensitivity curves of the compound eye of male and female A. corpulenta after dark adaptation of 60 min during the day.
Mean ±SE of six individuals.
Fig. 3 Spectral sensitivity curves of the compound eye of male and female A. corpulenta after dark adaptation of 60 min during the
night. Mean ±SE of five individuals.
Fig. 4 Spectral sensitivity curves of compound eye of A. corpulenta after light adaptation of 60 min. Mean ±SE of six individuals.
Fig. 5 Response-intensity curves of the compound eye of male and female A. corpulenta to white light after dark adaptation of 60 min
during the night. Mean ±SE of six individuals.
Fig. 6 Response-intensity curves of the compound eye of male and female A. corpulenta to white light after dark adaptation of 60 min
during the day. Mean ±SE of six individuals.
12
Fig. 1
1.0
female
male
0.9
0.8
Relative sentivity
0.7
0.6
0.5
0.4
0.3
0.2
0.1
320 340 360 380 400 420 440 460 480 500 520 540 560 580 600 620
Wavelength(nm)
Fig. 2
male
female
1.0
0.9
Relative sentivity
0.8
0.7
0.6
0.5
0.4
0.3
0.2
0.1
320 340 360 380 400 420 440 460 480 500 520 540 560 580 600 620
Wavelength(nm)
Fig. 3
13
1.0
male
female
0.9
0.8
Relative sentivity
0.7
0.6
0.5
0.4
0.3
0.2
0.1
320 340 360 380 400 420 440 460 480 500 520 540 560 580 600 620
Wavelength(nm)
Fig. 4
30
male
female
ERG reponse(mV)
25
20
15
10
5
0
0.0
0.5
1.0
1.5
2.0
2.5
3.0
Relative intensity(logI)
Fig. 5
14
3.5
4.0
4.5
5.0
30
male
female
ERG reponse(mV)
25
20
15
10
5
0
0.0
0.5
1.0
1.5
2.0
2.5
3.0
Relative intensity(logI)
Fig. 6
15
3.5
4.0
4.5
5.0