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. 2026 May 28;17(6):558. doi: 10.3390/insects17060558

Brightness Discrimination in the Crepuscular Moth Grapholita molesta (Busck, 1916) Under Dappled Light

Xiaofan Yang 1, Bao Li 2, Tongtong Huang 3, Guoshu Wei 2, Yafei Ge 2,*, Yanran Wan 2,*
Editor: David G James
PMCID: PMC13299949  PMID: 42355292

Simple Summary

The crepuscular moth Grapholita molesta is a major pest of fruit trees, and the females lay eggs on young leaves with a higher brightness or intensity around dusk. The light environment in orchards often varies between daylight and leaf shade, creating a dappled light condition. It is not known whether such dappled light affects brightness discrimination ability. In this study, we tested the preferences of G. molesta females between light-green (brighter brightness) and dark-green (darker brightness) stimuli, and between high- and low-light-intensity areas under uniform light, simulated dappled light, and complex dappled light conditions at three ecologically relevant illuminances (100, 1, and 0.01 lx). The results show that G. molesta females consistently preferred brighter green and higher-intensity stimuli in all light conditions, and dappled light did not reduce their brightness discrimination. These findings indicate that G. molesta females can reliably use brightness cues to choose oviposition sites, even in the challenging, heterogeneous light environments commonly found in orchards.

Keywords: Grapholita molesta, crepuscular moth, brightness discrimination, dappled light, oviposition preference

Abstract

The light environment in orchards is highly heterogeneous, characterized by dappled light caused by sunlight filtering through plant canopies, and poses a challenge for the color perception of insects. For the crepuscular moth Grapholita molesta, a global pest of fruit trees, females can visually discriminate between plants to locate young leaves with higher brightness or intensity for oviposition around dusk. However, whether dappled light affects this brightness discrimination performance remains unknown. In this study, we investigated the oviposition preference of G. molesta females between two brightness stimuli (light-green and dark-green), as well as among three different light intensities under uniform light, simulated dappled light, and complex dappled light conditions at three ecologically relevant illuminances (100, 1, and 0.01 lx). G. molesta females laid significantly more eggs on light-green than dark-green stimuli under all three light conditions at each illuminance. Similarly, females consistently preferred high-intensity over lower-intensity light areas across uniform, dappled, and complex dappled light conditions, with more than 66% oviposition frequency. Our findings show that the brightness and intensity discrimination ability of G. molesta is not affected by dappled light, which may enable moths to accurately locate suitable oviposition sites even when the orchard canopy creates a heterogeneous light distribution.

1. Introduction

Insects use color information for numerous tasks, including evaluating ambient light for phototaxis or navigation, detecting hosts for feeding or oviposition, and recognizing mates and predators for reproduction or survival [1,2]. In color perception, brightness is the most fundamental characteristic of a visual stimulus, describing the achromatic aspect associated with the intensity of perceived reflected or emitted light [3,4,5]. Although studies of insect color perception have generally focused on hue selectivity (i.e., the spectral distribution of reflected or emitted light), positive responses to brightness cues have been explicitly observed, particularly in some flower-visiting insects. For example, the diurnal species Eristalis tenax, Papilio xuthus and Macroglossum stellatarum all preferred the brighter of two shades with the same hue [6,7,8]. Given their higher signal-to-noise ratio, brightness cues may be particularly important under specific conditions, such as at night when light intensity is low. Accordingly, it has been reported that the crepuscular Manduca sexta and nocturnal Deilephila elpenor species exhibit a strong preference for higher brightness stimuli [5,9,10,11,12].

Natural illumination can vary significantly over the course of the day, between shaded and sunlit areas within a scene, and across different habitats such as forests and open fields [13,14]; these changes make reliable color perception particularly challenging. The insect visual system must therefore have mechanisms to compensate for differences in hue and brightness, i.e., color constancy or contrast constancy, to ensure that color information remains stable under changing illumination conditions [15,16]. For example, bees, butterflies, and hawkmoths can recognize learned target hues with high accuracy across varying illuminations [16,17,18,19,20]. In Drosophila, the luminance-sensitive pathway via the L3 neuron serves as a corrective signal that ensures contrast constancy, enabling accurate visual processing under rapidly changing light conditions [21]. Similarly, the bumblebee Bombus terrestris exhibits approximate color constancy and can discriminate between two similar hues differing in brightness (mauve and purple), although its accuracy is approximately 10% lower under simulated leaf shade than under daylight illumination [22].

The crepuscular oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae), is a major fruit-boring pest worldwide, causing significant economic damage to peach (Prunus persica L.), pear (Pyrus communis L.), apple (Malus domestica Borkh.), and other Rosaceae trees [23,24]. Gravid females prefer to lay eggs on the young leaves at the top of host plants around dusk [25,26]. While olfactory cues, specifically host volatiles, have been extensively documented as long-range orientation signals for distinguishing between host and non-host plants or among host species [27,28,29,30,31], visual cues become more important for perception in microhabitats. Our recent studies have shown that G. molesta females can visually discriminate oviposition sites in microhabitats even under dim light, with a preference for higher brightness and light intensity [26,32]. The light environment in orchards is heterogeneous; in particular, dappled light, which results from sunlight filtering through the branches and leaves of the plant canopy, contributes to this variability for much of the growing season. However, it remains unclear whether such dappled light environments affect brightness or intensity discrimination performance.

In this study, we investigated the oviposition preferences of G. molesta females between two brightness stimuli (light-green and dark-green) under simulated uniform, dappled, and complex dappled light conditions at three ecologically relevant illuminances (100, 1, and 0.01 lx, corresponding to twilight, late twilight, and moonlight). We also compared the preference for higher versus lower light intensities presented in choice tests.

2. Materials and Methods

2.1. Insects

A colony of G. molesta was established from wild-caught larvae in a peach orchard in Shunping, Hebei, China in August 2020. The colony was maintained in artificial climate chambers at 26 ± 1 °C, 70 ± 5% relative humidity (RH) under a 15 h:9 h light:dark cycle. Larvae were mass-reared on apples until they reached the third instar and were then transferred to an artificial diet [33] and adults were fed a 10% honey solution on cotton balls. Newly emerged adults were sexed and individually housed in glass tubes (1.8 cm diameter, 8 cm height) without access to food or water. Groups of fifteen 2-day-old females and males were paired 3–4 h before the onset of the scotophase. During the ensuing hours, copulating pairs were removed gently and separated 12 h later. Mated 3-day-old females were collected for oviposition preference experiments.

2.2. Experimental Setup and Stimuli

Experiments were conducted in a cuboid cardboard box (20 × 20 × 30 cm, Figure 1A) in a dark room at 26 ± 1 °C, 60% ± 5% RH. The floor of the box was covered by rough white paper to prevent undesired oviposition. The stimuli, hereafter called light-green (Ge170) and dark-green (Ge110), were represented by twelve rectangular papers (5 × 10 cm) arranged vertically in an alternating 4 × 3 grid on each side of the box, simulating young and old peach leaves, respectively. The light-green and dark-green rectangles were printed on white matte paper (100 g/m2) using an inkjet printer (L801, Seiko Epson Corp., Nagano, Japan) with custom RGB settings following Yang et al. (2020) [32]; their reflectance spectra (Figure 1D) show that they shared the same chromatic properties but differed in brightness (achromatic properties). Each paper was laminated with anti-reflection film to provide a smooth thigmotactic surface while preventing contamination from moth excreta. We simulated leaf-shade conditions with light-green and dark-green filters placed above the box. These filters were produced by printing on tracing paper (0.05 mm thick) with same RGB settings as those used for the paper rectangles.

Figure 1.

Figure 1

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Experimental setup and stimuli. (A) Schematic diagram of the experimental boxes used to test the oviposition preference of G. molesta. A white LED lamp was positioned 50 cm above the box, and light-green (Ge170) and dark-green (Ge110) colored stimuli were presented alternately on the box wall. (B) Pattern of simulated dappled light. Light-green and dark-green patches represent young and old leaf-shade light filters, while white patches indicate areas illuminated by LEDs. (C) The second layer of the complex dappled light pattern. (D) Reflectance spectra of colored papers. (E) Emission spectra of the illuminating white LEDs.

The experimental setup was illuminated from above (>50 cm) by a white LED lamp (18 W; Guangzhou Xinyuan Optoelectronic Technology Co., Ltd., Guangzhou, China). Corresponding to the light intensity which decreased with distance from the light source, the setup was vertically divided from top to bottom into three light-intensity areas without any physical barriers: a higher-intensity zone (top 1/3), a mid-intensity zone (middle 1/3), and a lower-intensity zone (bottom 1/3). The illumination was broad-spectrum with two emission peaks in the blue and green–yellow region (Figure 1E). Experiments were performed under three ecologically relevant illuminances: 100 lx (twilight), 1 lx (late twilight) and 0.01 lx (moonlight). Changes in illuminance do not affect the spectral distribution of LED-emitted light. Light intensity was measured at the top center of the experimental boxes using a radiometer (IL1700, International Light Research, Peabody, MA, USA) and adjusted by neutral density filters (Melles Griot, Rochester, NY, USA).

2.3. Experimental Procedure

To investigate whether dappled light affects brightness and intensity discrimination in G. molesta, we conducted three sets of experiments under illuminations of 100, 1 and 0.01 lx. First, oviposition preference was assessed under uniform lighting conditions. A large sheet of PVC plastic wrap was placed above the experimental box to ensure uniform illumination from the LEDs. Second, we created a dappled light environment by placing a leaf-shade filter above the box. The filter was arranged in a 4 × 4 alternating checkerboard patterns, consisting of eight colored squares (5 × 5 cm) and eight clear PVC-wrap squares (5 × 5 cm) (Figure 1B). This design allowed half of the box to be illuminated through the colored filters and the other half directly by LEDs, simulating the patchy light penetrating a leaf canopy. The colored squares were composed of two light-green and one dark-green patch [22]. Third, to introduce a more complex dappled light environment, we overlaid a second filter onto the checkerboard filter used in the second experiment. This additional filter was composed of alternating vertical stripes: two columns of light-green interleaved with dark-green stripes, and two columns of clear PVC wrap (Figure 1C). Each experiment was repeated 10 times.

All experiments began at 17:00, 3 h before the onset of the scotophase (when the moths start to oviposit). Ten 3-day-old mated females were introduced into the experimental boxes and allowed to lay eggs for 15 h overnight. The following morning, the females were removed, and the number of eggs laid on light-green and dark-green rectangles were recorded. When presenting stimuli to the moths, the anti-reflection laminating film was carefully cleaned with 75% alcohol. Females were tested only once.

2.4. Statistical Analysis

We used the Wilcoxon signed-rank test to compare the preference of G. molesta females between light-green and dark-green colors under different illuminations. Generalized linear models (GLMs) and Bonferroni post hoc tests were used to compare the preference across different light intensities, and to assess the differences in response to green brightness and light intensity under uniform light, dappled light and complex dappled light conditions. Females with a fecundity of fewer than 10 eggs were recorded but excluded from the statistical analysis. All statistical analyses were conducted using SPSS 27.0 (SPSS Inc., Chicago, IL, USA).

3. Results

3.1. Oviposition Responses to Green Brightness

G. molesta females exhibited a similar oviposition response to green brightness among uniform light, dappled light and complex dappled light conditions (Table 1, p > 0.05). Under all three light conditions, females significantly preferred to lay eggs on the light-green (Ge170) over the dark-green (Ge110) paper at illuminances of 100, 1 and 0.01 lx, with more than a 57% oviposition frequency (Figure 2 and Figure 3) (uniform light: Z100 lx = −2.547, Z1 lx = −2.803, Z0.01 lx = −2.366; dappled light: Z100 lx = −2.346, Z1 lx = −2.803, Z0.01 lx = −2.497; complex dappled light: Z100 lx = −2.547, Z1 lx = −2.805, Z0.01 lx = −2.803, all p < 0.05).

Table 1.

Influence of dappled light on brightness/intensity discrimination of G. molesta.

Light Intensity Oviposition Preference Tests F p-Value
100 lx Responses to green brightness 0.298 0.745
Responses to light intensity 0.006 0.994
1 lx Responses to green brightness 1.162 0.328
Responses to light intensity 0.000 1.000
0.01 lx Responses to green brightness 4.733 0.131
Responses to light intensity 0.000 1.000
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Figure 2.

Figure 2

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Oviposition preference of G. molesta in choices between light-green (Ge170) and dark-green (Ge110) colors under uniform light (A), dappled light (B) and complex dappled light (C). Data shown as mean ± SE, n = 10 experiments, 100 moths. Asterisks indicate significant differences (Wilcoxon signed-rank test; * p < 0.05; ** p < 0.01).

Figure 3.

Figure 3

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Number of eggs laid by G. molesta females on light-green and dark-green rectangles in higher-intensity (HI), mid-intensity (MI) and lower-intensity (LI) zones under illuminances of 100, 1 and 0.01 lx. (A–C) Oviposition preferences were tested under uniform light (A), dappled light (B) and complex dappled light (C). Each point represents the number of eggs laid by a single female. Orange, deep blue and green represent the mapping of each point onto the X-, Y-, and Z-axes, respectively.

3.2. Oviposition Responses to Light Intensity

Under uniform light, G. molesta females consistently exhibited a strong preference to lay eggs in the higher-intensity (HI) areas, with oviposition frequencies of 66.16%, 85.39% and 78.10% at illuminances of 100, 1 and 0.01 lx, respectively, and fewer eggs were laid in mid-intensity (MI) and lower-intensity (LI) areas (100 lx: F = 67.482, p < 0.001; 1 lx: F = 455.549, p < 0.001; 0.01 lx: F = 375.394, p < 0.001). Similarly, females significantly preferred HI areas under both dappled light and complex dappled light across all tested illuminances (Figure 3 and Figure 4, p < 0.001). These results suggest that the intensity discrimination of G. molesta is not affected by dappled light (Table 1, p > 0.05).

Figure 4.

Figure 4

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Oviposition preference of G. molesta in choices among higher-intensity (HI), mid-intensity (MI) and lower-intensity (LI) conditions under uniform light (A), dappled light (B) and complex dappled light (C). Data shown as mean ± SE, n = 10 experiments, 100 moths. Different letters above bars indicate significant differences at the p < 0.05 level (GLM followed by Bonferroni test).

4. Discussion

As a moth moves through orchards in search of leaves or fruits for oviposition, it experiences different illuminations, from bright daylight to patchy leaf shade. Here, we investigated whether such dappled light environments affect brightness discrimination in the crepuscular moth G. molesta. Our results demonstrate that G. molesta females consistently preferred light-green over dark-green stimuli, and higher-intensity over lower-intensity light areas, across uniform, dappled, and complex dappled light conditions under the ecologically relevant illuminances tested (100, 1, and 0.01 lx). These findings suggest that brightness and intensity discrimination in G. molesta remains stable across heterogeneous light environments, and so they may be able to recognize oviposition sites with high accuracy in orchards where dappled light prevails for much of the growing season. To distinguish differences in brightness or light intensity under changing illumination conditions, G. molesta requires a highly sensitive visual system. Indeed, this crepuscular moth possesses superposition compound eyes, which are characterized by high light capture ability due to the presence of a clear zone separating crystalline cones from the rhabdom layer [34]. This structural adaptation improves visual reliability. Potentially, this reliability can be further improved by neural summation of light in space and time, which substantially increases contrast sensitivity and visual information rate, thereby supporting stable visual processing under rapidly changing light conditions [35].

Color or contrast constancy, the ability to perceive hue or brightness as stable under changing illumination, requires the visual system to compensate for illumination changes [15,16,20]. Here, the failure of dappled light to alter oviposition preferences for brightness and intensity in G. molesta suggests that females may employ luminance-based contrast constancy mechanisms, similar to those described in Drosophila [21]. Briefly, luminance-based contrast constancy is the ability of an insect’s visual system to detect a potential plant or predator by the luminance of the object against the luminance of the background. Alternatively, the moths might have evolved effective visual adaptation mechanisms to integrate brightness information over space or time, averaging out local variations introduced by leaf-shade patterns. Such strategies would be adaptive for a crepuscular species active during periods of rapid and patchy light changes. Future studies could investigate the luminance processing pathways and retinal mechanisms of G. molesta to elucidate the mechanistic basis of its robust brightness discrimination under variable light.

In contrast to our findings, Arnold and Chittka (2012) reported that bumblebees B. terrestris showed reduced color discrimination accuracy under simulated leaf shade, particularly when distinguishing between similar hues that differed primarily in brightness [22]. This discrepancy may reflect differences in visual ecology: diurnal bees rely heavily on chromatic cues for flower discrimination, whereas crepuscular G. molesta may depend more on achromatic (brightness) cues, which are less affected by spectral shifts in shade and can be detected with a higher signal-to-noise ratio in dim light [5,36].

The preference for brighter and higher-intensity stimuli observed in G. molesta across all experimental conditions aligns with our previous findings that this species relies on brightness or intensity cues for oviposition in dim light [26,32] and is consistent with the visual behavior of other crepuscular and nocturnal lepidopterans. For example, the crepuscular M. sexta and nocturnal D. elpenor both show strong innate preferences for higher-intensity stimuli [5,10,11,12]. This study extends these findings by demonstrating that this preference persists even under dappled light, thus allowing G. molesta females to accurately locate oviposition sites even when the orchard canopy creates patchy light distribution. This adaptation may enhance offspring survival, as young leaves at the top of host plants provide optimal nutritional conditions for larval development [37].

Notably, our experimental females were socially naive, reared in isolation from eclosion to oviposition. Social isolation can alter insect oviposition behavior: isolated females of flies, cockroaches, and beetles often lay fewer eggs or show altered host preferences compared to grouped conspecifics [38,39,40]. Future studies should test socially experienced females to validate whether social cues modulate brightness-based oviposition choices in G. molesta. While our study isolated visual cues to test brightness discrimination, insects often use both visual and olfactory cues in nature. Under field conditions, brightness preferences likely interact with olfactory cues: a bright leaf may be rejected if it emits repellent volatiles, while a dim leaf may be accepted if it releases attractive volatiles. Our findings provide a foundational understanding of visual preference, but field studies integrating both visual and chemical cues are needed to fully explain oviposition site selection in G. molesta.

In conclusion, our study provides strong evidence that G. molesta females maintain reliable brightness discrimination across uniform, dappled, and complex dappled light environments, at illuminances ranging from twilight to moonlight. These findings advance our understanding of insect visual adaptation to variable light environments and may have implications for developing visual-based pest management strategies, such as optimizing the brightness contrast of traps or repellent surfaces. Future research into the mechanistic basis of brightness discrimination in G. molesta will further elucidate the evolutionary and physiological adaptations of crepuscular insect visual systems to low and heterogeneous light.

Acknowledgments

We would like to thank Fan Zhang and Liyang Wang for rearing the insects.

Author Contributions

Conceptualization, X.Y. and Y.W.; methodology, X.Y. and Y.G.; software, B.L.; validation, B.L. and T.H.; formal analysis, X.Y.; investigation, X.Y. and B.L.; resources, X.Y. and Y.W.; data curation, Y.G.; writing—original draft preparation, X.Y.; writing—review and editing, Y.G. and Y.W.; visualization, G.W.; supervision, G.W. and Y.W.; project administration, X.Y.; funding acquisition, X.Y. All authors have read and agreed to the published version of the manuscript.

Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding authors.

Conflicts of Interest

The authors declare no conflicts of interest.

Funding Statement

This research was funded by the National Natural Science Foundation of China (32402368) and the Natural Science Foundation of Hebei Province (C2024301061).

Footnotes

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Data Availability Statement

The original contributions presented in this study are included in the article. Further inquiries can be directed to the corresponding authors.

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