Multimodal stimulation of Colorado potato beetle reveals modulation of pheromone response by yellow light

Orientation of insects to host plants and conspecifics is the result of detection and integration of chemical and physical cues present in the environment. Sensory organs have evolved to be sensitive to important signals, providing neural input for higher order multimodal processing and behavioral output. Here we report experiments to determine decisions made by Colorado potato beetle (CPB), Leptinotarsa decemlineata, in response to isolated stimuli and multimodal combinations of signals on a locomotion compensator. Our results show that in complete darkness and in the absence of other stimuli, pheromonal stimulation increases attraction behavior of CPB as measured in oriented displacement and walking speed. However, orientation to the pheromone is abolished when presented with the alternative stimulation of a low intensity yellow light in a dark environment. The ability of the pheromone to stimulate these diurnal beetles in the dark in the absence of other stimuli is an unexpected but interesting observation. The predominance of the phototactic response over that to pheromone when low intensity lights were offered as choices seems to confirm the diurnal nature of the insect. The biological significance of the response to pheromone in the dark is unclear. The phototactic response will play a key role in elucidating multimodal stimulation in the host-finding process of CPB, and perhaps other insects. Such information might be exploited in the design of applications to attract and trap CPB for survey or control purposes and other insect pests using similar orientation mechanisms.     

Impact of Host Plant Connectivity,Crop Border and Patch Size on Adult Colorado Potato Beetle Retention

Tagged Colorado potato beetles (CPB), Leptinotarsa decemlineata (Say), were released on potato plants, Solanum tuberosum L., and tracked using a portable harmonic radar system to determine the impact of host plant spatial distribution on the tendency of the pest to remain on the colonized host plant or patch. Results confirmed the long residency time on the host plant and showed that close connection of the plant to neighboring plants hastened dispersal between plants. Tracking walking CPB for over 6 h in small potato plots revealed that all types of mixed borders tested (potato/bare ground, potato/timothy and potato/woodland) acted as a strong barrier and retained beetles within the patch. In another experiment in potato patches surrounded by bare ground borders, tracked walking CPB displayed similar behaviour for up to four days. The distribution of turning angles in the CPB walking paths was not uniform and corresponded to beetles following the edge rows of potato patches in response to the crop border barrier or reversing their direction as they reached the end of a row and therefore a border. Patch size had no or little effect on beetle retention in the patch. The relative distribution of counts of tagged beetles detected among small (16 m²), medium (64 m²) and large size (256 m²) patches of potato four days after initial release remained similar to that of numbers released. Even though mixed crop borders were a strong barrier to walking CPB emigrating from potato patches, the departure rate of beetles over time was high. Results suggest that the effect of mixed borders is largely limited to dispersal by walking and does not apply to beetles leaving host patches by flight. The manipulation of crop borders and patch size seem to have limited potential for the management of CPB emigrating from potato fields.     

Spectral preferences of Lycorma delicatula (Hemiptera: Fulgoridae)

Many insects prefer lights with certain spectral properties, and such preferences may be associated with behavioral contexts such as mating, host finding or dispersal. Lycorma delicatula (Hemiptera: Fulgoridae) is a newly invasive species in Korea and is spreading rapidly. It is diurnal and may rely on visual cues for orientation. We conducted a series of three phototaxis experiments to understand spectral preferences in L. delicatula: light/dark choice, UV/white light choice, and color preference experiments. Nymphs of the third and final stages as well as adults were used for these experiments. In the light/dark choice, the orientation of L. delicatula was bimodal between the white light and darkness, for all stages and both sexes. In a choice of UV (395–410 nm wavelengths) vs. white light, L. delicatula of both sexes and all stages preferred the UV light. In the color preference experiment where insects had a choice of four colors in a circular arena, L. delicatula stayed significantly longer in the blue light than in white, yellow or green lights. Overall, nymphs and adults of L. delicatula oriented toward lights with shorter wavelengths, and this orientation was consistent throughout all stages, regardless of sex. It is necessary to investigate the behavioral contexts under which L. delicatula prefers the UV and blue lights.     

Visual orientation of the black fungus gnat,Bradysia difformis,explored using LEDs

Fungus gnats occur worldwide with more than 1 700 described species. They can cause serious damages on ornamentals, crop plants, and edible mushrooms, and are considered to be a serious pest in the last years. Bradysia difformisFrey (Diptera: Sciaridae) represents a common species in Europe. Usually, yellow sticky traps are used for monitoring and control in greenhouses and fluorescent tube‐based light traps are additionally applied for control in mushroom cultivation. The importance of such visual trapping measures for efficient monitoring or alternative control increases in biological and integrated plant protection. However, detailed color preferences of fungus gnats are mostly unknown. We studied the visual orientation of B. difformis with light‐emitting diodes (LEDs) in a broad range of peak wavelengths from 371 nm (ultraviolet, UV) to 619 nm (amber). We determined attractive wavelengths in consecutive choice experiments in daylight and darkness. Highest numbers of adult B. difformis were attracted to UV radiation (382 nm) followed by green‐yellow light (532–592 nm). The responses to UV and the green‐yellow range were relatively unspecific and mostly independent from intensity. Combination of UV and yellow LEDs improved trapping efficacy compared to a single UV or yellow LED trap, as well as compared to a common yellow sticky trap. When both wavelengths were compared to a black surface to increase contrasts, the black surface was preferred over yellow, but was less attractive than UV. Thus, B. difformis displays two, probably wavelength‐specific, behaviors to UV radiation and green‐yellow light, with UV being the most attractive stimulus. These behaviors might be directly related to underlying photoreceptors, suggesting dichromatic vision in B. difformis.     

Which wavelength does the cigarette beetle, Lasioderma serricorne (Coleoptera: Anobiidae), prefer? Electrophysiological and behavioral studies using light-emitting diodes (LEDs)

The cigarette beetle, Lasioderma serricorne (Fabricius), is an important pest insect that consumes a variety of dry foods. It is known that UV light traps attract this species. However, less attention has been paid to its preferred wavelength. First, we investigated the spectral sensitivity of the compound eye. Next, we compared the attraction efficiency of LEDs of different colors (wavelengths). Our results showed that ultraviolet (UV, 375 nm) and blue (470 nm) LEDs attracted the most cigarette beetles of both sexes, irrespective of mating or oviposition status, although the UV LED consistently tended to attract the most beetles. Although the primary sensitivity peak of the compound eye was 520 nm, the green LED (520 nm) scarcely attracted beetles. Although the reason for the difference between the peaks in spectral sensitivity and attraction of beetles awaits further studies, whether UV and/or blue LEDs is more effective as a practical light trap for controlling L. serricorne beetle is discussed in this study.     

Contribution of habitat type to residency and dispersal choices by overwintered and summer adult Colorado potato beetles

The walking and flight dispersal of marked overwintered and summer Colorado potato beetles (CPB), Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), released in field box‐plots was monitored simultaneously in six habitats over a period of 4 days. The emigration out of plots by walking beetles was calculated from the catch in linear pitfall traps completely surrounding each box‐plot and emigration flight was estimated from the number of beetles missing from the plot or captured by the trap. Overwintered beetles dispersed sooner after release than summer beetles. Overall, the mean number of beetles retained by the habitat was significantly higher in the host habitat (potato) than in any non‐host habitat tested (soybean, pasture, bare ground, water, woodland). Unexpectedly, there was no or little difference in overall beetle retention between non‐host habitats except for higher retention in the water habitat. No difference in the ratio of flight over walking could be detected by the study between overwintered and summer CPB except in the water and woodland habitats. Twenty‐four hours after release, the highest ratios were obtained in the water and woodland habitats and the lowest in the bare‐ground habitat, but ratios were similar for all habitats, except water, after 96 h. As a population, under these experimental conditions, 96 h after release, it seems that CPB displayed a slight preference for flight over walking, with walking as a default mode. A fed and starved pre‐release treatment had no effect on dispersal rates or mode of dispersal. Essentially, our results showed that over a 96‐h period, northeastern North American CPB emigrated at similar rates from the various non‐host habitats encountered, except for water, using walking as much as flight. The host habitat retained CPB significantly longer than non‐host habitats but with a mode of dispersal ratio similar to that in non‐host habitats. The impact on dispersal of the various habitats encountered by CPB in the agro‐ecosystem was less important than expected suggesting that the interaction of environmental parameters is likely to have the most significant impact in determining dispersal rates and dispersal modes.     

Photoperiodic and light spectral conditions which inhibit circulating concentrations of thyroxine in the male hamster

Adult male Syrian hamsters were exposed daily for 12 weeks to 11 h/day of cool white fluorescent light (350 +/- 50 microW/cm2) followed by an additional 3 h of near ultraviolet (339-317 nm), blue (435-500 nm), green (515-550 nm), yellow (558-636 nm) or red (653-668 nm) light at an irradiance of 0.2 microW/cm2 or to total darkness. Animals exposed to the wavelengths between 558-668 nm (yellow or red half peak bandwidths) or those receiving a total of 13 h of darkness/day had suppressed circulating levels of thyroxine (T4), a depressed free T4 index (FT4I) and a higher T3/T4 ratio compared to animals receiving a total of 14 h of white light (350 +/- 50 microW/cm2). These results suggest that specific wavelengths of light can affect the neuroendocrine-thyroid axis.     

Role of visual stimuli in host and mate location of the Colorado potato beetle

Visual responses of adult male and female Colorado potato beetles CPB, Leptinotarsa decemlineata (Say) (Coleoptera, Chrysomelidae) to 10 × 20 cm coloured paper‐boards, beetle‐sized coloured beads or dead CPB, and to combinations of these in laboratory experiments in an arena of 52 cm diameter were investigated. In addition, visual discriminatory abilities of males exposed to female‐specific substances were also measured. The paths of the beetles were recorded on videotape by a computer‐aided motion detection system. Parameters of tracks were calculated and the tape‐recorded paths were further processed for behavioural analyses and for obtaining mean angular directions by circular statistics. Both sexes showed a strong preference for yellow‐black striped, yellow and light‐green boards and, to a lesser degree, were also attracted to yellow‐green, dark‐green, red and white boards, to beetle‐sized beads coloured either black or yellow‐black striped, and to some combinations of these. Males exposed to female substances immediately prior to entering the arena performed significantly more successful approaches to objects than did males of the control group. These results corroborate field observations on males' mate‐searching behaviour and increase our understanding of natural host‐ and mate‐finding processes.     

Light intensity and wavelength influence development, reproduction and locomotor activity in the predatory flower bug Orius sauteri (Poppius) (Hemiptera: Anthocoridae)

Light wavelength and intensity are physical factors that can affect arthropod development and reproduction. The present study examined the development, reproduction and locomotor activity of the predatory flower bug, Orius sauteri (Poppius) (Hemiptera: Anthocoridae), under five light intensities (1000, 2000, 3000, 4000 and 5000 lux) and five wavelengths [red (678.5 nm), green (620.0 nm), yellow (581.7 nm), blue (478. 1 nm) and white (all wavelengths)] at constant temperature (25 °C) and RH (70 %). The duration of nymphal development was extended at lower light intensities, primarily due to effects on the first three instars. Under white, yellow and green light, O. sauteri completed development in 18.0 days, but blue light extended development by 3.2 days and red light extended it by 7.4 days. Although lower light intensities extended the preoviposition period and reduced fecundity, they improved egg fertility. Both red and blue light negatively affected preoviposition period, fecundity and egg fertility. Whereas adult female mean walking speed over a five min period was reduced at lower light intensities, longer wavelengths (yellow and red) increased it, ostensibly reflecting an avoidance response. The respiration quotient of adult O. sauteri females was also elevated under red light conditions. These findings are informative for optimizing O. sauteri mass-rearing procedures and maximizing its efficacy as a biological control agent in greenhouse cultures.     

Flight take-off performance of Colorado potato beetle in relation to potato phenology

The flight take-off frequency of adult Colorado potato beetles, Leptinotarsa decemlineata (Say), from potato plants, Solanum tuberosum L. 'Red Pontiac' at the bloom stage of development was 2.2-2.5-fold that of Colorado potato beetle from plants at the vegetative stage. Tests were conducted in a flight chamber over a period of 3 h. Prefeeding Colorado potato beetles for 48 h on potato plants at the bloom or at the vegetative stage before placing them into the flight chamber resulted in the same significantly higher flight take-off frequency from potato plants at the bloom stage than from plants at the vegetative stage. These results demonstrate that the factor in potato plants in bloom that stimulates the flight take-off of the Colorado potato beetle is independent of the feeding history of the beetles and begins acting only when the beetles are in the presence of the plant. According to these results, the dispersal of adult Colorado potato beetles from potato fields in bloom to younger potato fields with plants at the vegetative stage, previously reported in the literature, is at least partly explained by the effect of plant phenology on the frequency of flight take-off. Results confirm the value of planting potato fields of similar phenology over as wide an area as possible to reduce Colorado potato beetle dispersal between fields. Results also imply that staggering the planting dates of conventional potato refuge areas near Colorado potato beetle transgenic or conventionally resistant potato fields is a sound management practice, because it promotes the movement of wild beetles over to the adjacent younger resistant crops.     

Role of visual stimuli in host and mate location of the Colorado potato beetle

Visual responses of adult male and female Colorado potato beetles CPB, Leptinotarsa decemlineata (Say) (Coleoptera, Chrysomelidae) to 10×20 cm coloured paper-boards, beetle-sized coloured beads or dead CPB, and to combinations of these in laboratory experiments in an arena of 52 cm diameter were investigated. In addition, visual discriminatory abilities of males exposed to female-specific substances were also measured. The paths of the beetles were recorded on videotape by a computer-aided motion detection system. Parameters of tracks were calculated and the tape-recorded paths were further processed for behavioural analyses and for obtaining mean angular directions by circular statistics. Both sexes showed a strong preference for yellow-black striped, yellow and light-green boards and, to a lesser degree, were also attracted to yellow-green, dark-green, red and white boards, to beetle-sized beads coloured either black or yellow-black striped, and to some combinations of these. Males exposed to female substances immediately prior to entering the arena performed significantly more successful approaches to objects than did males of the control group. These results corroborate field observations on males' mate-searching behaviour and increase our understanding of natural host- and mate-finding processes.     

Can ultraviolet cues function as aposematic signals?

The fact that birds are sensitive to ultraviolet light (UV,320-400 nm) has been largely ignored by previous studies ofaposematism. Therefore, in the present article we investigatedwhether great tits preferred ultraviolet-reflecting colorscompared to colors without UV reflection and whether UV cuesalone could function as aposematic signals. We were able to manipulate prey visibility in UV light by changing the UV reflectanceof prey items as well as altering the lighting conditions.In order to perform a preference experiment we used three pairsof colors (green+UV vs. green, gray+UV vs. gray, yellow+UVvs. yellow) on a black background. The birds ate both UV typesequally for all three colors. Thus, there was no avoidance of the UV-reflecting prey. Next we tested the possibility thatUV reflection may affect avoidance learning. We used eithergreen or green+UV as a signal for unpalatability. In this set-upthe difference in UV did not allow avoidance learning to occur.Our experiment with great tits does not support the hypothesisthat UV cues alone might work effectively as aposematic signals.     

不同波长光照对草履虫增殖的影响及其感光蛋白基因克隆的初步探究

目的研究不同波长光照对草履虫增殖的影响,克隆草履虫感光蛋白基因.方法以双小核草履虫 Paramecium aureli为研究对象,分别置于黄色光(578～592 nm)、蓝色光(446～464 nm)、红色光(620～760 nm)、白光和自然光下,每隔1 h随机抽样法显微观察并计数;RT-PCR克隆草履虫感光蛋白基因.结果不同波长光的照射下,与自然光比较,第1天黄光组草履虫增殖显著上升(P<0.01),蓝光组和红光组草履虫增殖受到不同程度的抑制(P<0.01, P<0.05),白光组无明显差异(P>0.05);第2天,蓝光组、红光组和白光组草履虫增殖仍受到抑制(P<0.01),黄光组作用不显著(P>0.05).黄光组和白光组草履虫总RNA作为模板,克隆出大小约500 bp的rhodopsin-like基因cDNA片段, 5个不同光照组均克隆出大小约195 bp的Long wave sensitive opsin-like 基因cDNA片段.结论黄色光显著地促进草履虫增殖,蓝色光和红色光抑制其增殖;黄光和白光能诱导草履虫rhodopsin-like基因表达;Long wave sensitive opsin-like 基因在草履虫有表达.     

Colour preference and potato cultivar oviposition choice by onion thrips,Thrips tabaci (Thysanoptera: Thripidae)

Thrips tabaci (Lindeman) is an important polyphagous pest, and vector of Tomato spotted wilt virus responsible for sporadic, but devastating epidemics in potato. T. tabaci shows significant preferential differences between potato cultivars that may be important for field resistance. To investigate the role of visual cues in host choice we tested colour preference using two-choice assays. Thrips tabaci showed a significant preference for mid-green over red, blue, and white coloured cards, a preference for both light-green and mid-green over dark-green, and light-green over yellow, but no preferential difference between mid-green and yellow, and between light-green and mid-green. Analysis of the spectral reflectance of potato cultivars differing in thrips preference, revealed significant differences within the 400–700 nm wavelengths. In most cases cultivars preferred by T. tabaci had lighter green foliage and higher reflectance at 552 nm, and thus colour preference may be important for host selection. Oviposition choice of T. tabaci for potato cultivars was determined from counts of larvae and unhatched eggs from leaf disks under choice and no-choice conditions. In contrast to the colour choice experiments, onion thrips showed oviposition preference for cultivars with darker green foliage and lower reflectance at 552 nm.     

多种植物挥发物及马铃薯甲虫聚集素对马铃薯甲虫的引诱作用

马铃薯甲虫Leptinotarsa decemlineata (Say)是马铃薯上一种最主要的毁灭性害虫。由于其严重的抗药性, 急需寻找一种替代杀虫剂的防治方法。本实验从开发马铃薯甲虫引诱剂的目的出发,用“Y”型嗅觉仪测定了马铃薯甲虫对来自植物的7种挥发物单体、8个挥发物混合物配方以及马铃薯甲虫聚集素的行为反应,并进一步进行了田间诱集试验。室内生测结果表明,2-苯乙醇对马铃薯甲虫雌雄虫都有明显的引诱作用。芳樟醇+水杨酸甲酯+顺乙酸-3-己烯酯的混合物对马铃薯甲虫雄虫有很强的引诱作用(81.67%), 但是对雌虫引诱作用不明显(63.33%)。在其中加入马铃薯甲虫聚集素不但提高了对雄虫的引诱率（88.33%）,还消除了之前的雌雄性别间反应差异现象,使对雌虫的引诱率达到了83.33%。田间诱集结果显示,芳樟醇+水杨酸甲酯+顺乙酸-3-己烯酯+马铃薯甲虫聚集素在所研究的引诱剂配方中引诱效果最好。从而为马铃薯甲虫发生的预测预报和诱杀技术提供了研究基础。     

Host plant choice experiments of Colorado potato beetle (Coleoptera: Chrysomelidae) in Virginia

Field and laboratory-choice experiments were conducted to understand aspects of host plant orientation by the Colorado potato beetle, Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae), in Virginia. In laboratory bioassays, L. decemlineata oriented to volatiles emitted by potato, Solanum tuberosum L., foliage over both tomato, Lycopersicon esculentum L., and eggplant, Solanum melongena L., foliage, and eggplant over tomato foliage, all of which had been mechanically damaged. Field choice tests revealed more L. decemlineata adults, larvae, and egg masses on eggplant than on tomato. In other experiments, counts of live L. decemlineata on untreated paired plants and counts of dead beetles on imidacloprid-treated plants did not differ between potato and eggplant. L. decemlineata was significantly attracted to eggplant over both tomato and pepper. To determine whether feeding adults affected orientation to host plants, an imidacloprid-treated eggplant or potato plant was paired with an untreated eggplant or potato plant covered in a mesh bag containing two adult male beetles. Significantly more adults were attracted to eggplant with feeding male beetles paired with another eggplant than any other treatment combination. These results indicate that the presence of male L. decemlineata on plants affects host plant orientation and suggests that the male-produced aggregation pheromone may be involved.     

Ultraviolet and violet light: attractive orientation cues for the Indian meal moth, Plodia interpunctella

The Indian meal moth (IMM), Plodia interpunctella (Hübner) (Lepidoptera: Pyralidae), engages in long-distance or foraging flights in the twilight hours of the scotophase when blue light dominates the irradiance spectrum of the sky. We tested the hypothesis that IMM uses wavelengths of visible blue/violet light as orientation cues that trigger phototactic responses. In four-choice laboratory experiments, blue light (400–475 nm) was significantly more effective than green (475–600 nm), orange (575–700 nm), or red (590–800 nm) light in attracting males and mated females. In subsequent experiments that tested light emitting diodes (LEDs) emitting peak wavelengths in the blue/violet-light range, the 405-nm 'violet' LED was significantly more effective than the 435-, 450-, or 470-nm 'blue' LED in attracting males as well as virgin and mated females. In electroretinogram recordings, the 405-nm wavelength elicited significantly stronger receptor potentials from female and male eyes than the 350-nm (UV) wavelength, and in a behavioral experiment it significantly enhanced the known attractiveness of UV light. Equal attraction of IMMs to 405-nm LEDs at 600–700 µW/cm² with or without UV light, and significantly stronger attraction to a 405-nm LED than to a 350-nm LED at maximum light intensities, suggest that the deployment of violet instead of UV light could become one of several management tactics for control of IMMs.     

Role of Vision and Mechanoreception in Bed Bug,Cimex lectularius L. Behavior

The role of olfactory cues such as carbon dioxide, pheromones, and kairomones in bed bug, Cimex lectularius L. behavior has been demonstrated. However, the role of vision and mechanoreception in bed bug behavior is poorly understood. We investigated bed bug vision by determining their responses to different colors, vertical objects, and their ability to detect colors and vertical objects under low and complete dark conditions. Results show black and red paper harborages are preferred compared to yellow, green, blue, and white harborages. A bed bug trapping device with a black or red exterior surface was significantly more attractive to bed bugs than that with a white exterior surface. Bed bugs exhibited strong orientation behavior toward vertical objects. The height (15 vs. 30 cm tall) and color (brown vs. black) of the vertical object had no significant effect on orientation behavior of bed bugs. Bed bugs could differentiate color and detect vertical objects at very low background light conditions, but not in complete darkness. Bed bug preference to different substrate textures (mechanoreception) was also explored. Bed bugs preferred dyed tape compared to painted tape, textured painted plastic, and felt. These results revealed that substrate color, presence of vertical objects, and substrate texture affect host-seeking and harborage-searching behavior of bed bugs. Bed bugs may use a combination of vision, mechanoreception, and chemoreception to locate hosts and seek harborages.     

Evaluating light attraction to increase trap efficiency for Tribolium castaneum (Coleoptera: Tenebrionidae)

The red flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), is a major coleopteran pest in flour mills and storage facilities. An aggregation pheromone has been identified for this pest; however, the pheromone is of limited value for population monitoring. To develop more efficient methods to monitor this pest, experiments were conducted to determine whether light functioned as an attractant for the red flour beetle. Light-emitting diodes (LEDs) of various wavelengths were examined as light sources because they produce bright, narrow light spectra. A comparison of responses to light spectra across the visible and UV regions of the electromagnetic spectrum indicated that the beetle was most attracted to near UV LED at a 390 nm dominant wavelength. The use of LEDs in competitive laboratory experiments resulted in a 20% capture of released beetles, compared with a 1% capture with the aggregation pheromone alone. Even more beetles were captured with a combination of LEDs and commercially available chemical lures in traps. LEDs can easily be added onto existing trap designs or new traps can be designed to take full advantage of positive phototaxis.     

The Influence of Light Wavelength on Phase-Dependent Responsiveness of the Photoperiodic Clock in Migratory Blackheaded Bunting

We investigated the phase-dependent effects of light wavelength on photoperiodic clock in the migratory blackheaded bunting. Two experiments were performed, employing a skeleton paradigm (6 hours light : 6 hours darkness : 1 hour light : 11 hours darkness; 6L : 6D : 1L : 11D) at 37 ± 2 lux intensity. In the experiment 1, both 6 and 1 h light pulses were given at the same wavelength, 500 nm (green) or 650 nm (red). A group exposed to both pulses of white light served as control. In the experi-ment 2, the two light pulses were given at two different wavelengths, 6 h at 500 nm (green) and 1 h at 640 nm (red) in one group or vice-versa in the other. There was almost no photoinduction when both light pulses in experiment 1, or 1 h light pulse in experiment 2, were green. On the other hand, birds fattened and testes recrudesced when both the light pulses in experiment 1, or 1 h light pulse in experiment 2, were red. Birds receiving both pulses of white light in experiment 1 showed an intermediate response. Taken together, these results indicate that the photoperiodic clock in buntings is differentially responsive at its various circadian phases to different light wavelengths.