Spectral preference and temporal modulation of photic orientation by Colorado potato beetle on a servosphere

The Colorado potato beetle (CPB), Leptinotarsa decemlineata (Say) (Coleoptera: Chrysomelidae), is a diurnal insect that strongly relies on visual cues to guide its walk. In the present study, we investigated the orientation behavior of non‐diapausing walking CPB in response to emissive colors produced by light emitting diodes (LEDs) in a dual choice arena adapted to a servosphere, where the only illumination available came from the photo‐stimuli. Our results demonstrate that CPB show positive phototactic behavior when stimulated with different wavelengths of light; they preferred to orient towards white (420–775 nm), ultraviolet (UV) (351 nm), blue (472 nm), green (570 nm), yellow (585 nm), orange (590 nm), and red (660 nm) over darkness when both alternatives were offered, but no orientation responses were elicited by infrared (940 nm). Both males and females preferred yellow and green over other colors, but did not show any particular preference between them, thus correlating with their preference for wavelengths reflected by vegetation. Sexual differences were noted in that male CPB preferred white over either red or UV, whereas female CPB did not show any preferences when offered these colors. Female CPB preferred UV and blue over red, whereas males showed no preferences when these colors were offered. Colorado potato beetles turned at higher angles and performed more tortuous walks in complete darkness and when infrared vs. darkness were offered compared with the rest of the colored lights. Both sexes preferred continuous over pulsed yellow light. Colorado potato beetles subjected to pulsed yellow light showed a temporal alteration of their walking performance by walking less, slower, and turning at a higher rate. The results are discussed with regard to the role of color in the CPB attraction to host plants and conspecifics as well as the role of intermittent photic stimuli in their orientation behavior. The information provided here provides a basis for the improvement of trapping devices for detection and survey of incipient or invasive CPB, and development of alternate control strategies for this important pest of potatoes and other solanaceous crops.     

昆虫趋光性研究的回顾

利用昆虫的趋光性开展农林卫生害虫诱杀,一直是害虫绿色防控的组成部分。本文综述了昆虫的多种趋光性行为,昆虫趋光性机制—罗盘理论、马赫带效应理论和开放空间理论,对在物理、生物环境条件、光和光源属性条件下昆虫趋光的生态适应性,以及对昆虫抵达光源的位置和移动轨迹进行了讨论。     

Aggregation in the haematophagous bug Triatoma infestans: a novel assembling factor

Abstract. The response of Triatoma infestans (Klug) to odours left behind by walking insects was analysed. The insects aggregated on papers marked with putative chemical footprints in the absence of other assembling chemical signals. Contact of the receptor insect with the treated surface was necessary to elicit the response because the signal was not able to attract insects, only to arrest them on marked surfaces. These requirements contrast with other chemical aggregation signals of triatomines that attract the insect and involve olfactory responses. By washing the cuticle of the insects with hexane and impregnating papers with the extract, a response was evoked in test insects similar to that elicited by papers on which insects had walked. The biological role of this novel aggregation factor is discussed in relation to other assembling signals of different origin.     

Orientation of flying male Anobium punctatum (Coleoptera: Anobiidae) to sex pheromone: separating effects of visual stimuli and physical barriers to wind

Abstract. Flying male woodworm, Anobium punctatum , are able to locate a source of female pheromone offered as a point source but land more quickly if the same odour source is presented on a model 'tree' (cylinder). We tested the response to a two-dimensional black 'tree' marked as a silhouette on the upwind screen of the wind tunnel, with or without a hidden upwind barrier to the wind. The beetles responded equally to pheromone presented on a visual silhouette with wind barrier and to the hidden upwind wind-barrier alone, suggesting that the wind-barrier itself has a significant effect on insect orientation and landing: visual stimuli do not add to the landing response if the barrier is present. However, without the barrier upwind, landing was faster when the visual stimulus was present than without. This is the third forest-living species to be shown to respond in this way, which suggests this phenomenon may be more widespread. The possibility that this phenomenon is present in other, non-forest, phytophagous insects, their predators and parasitoids is discussed.     

Phototactic behaviour of Pachyneuron aphidis (Hymenoptera: Pteromalidae) – hyperparasitoid of Myzus persicae (Hemiptera: Aphidiae)

We investigated the spectral sensitivity and response to light intensity of Pachyneuron aphidis (Hymenoptera: Pteromalidae), which is a common hyperparasitoid of Aphidius gifuensis (Hymenoptera: Braconidae), a key natural enemy of the green peach aphid, Myzus persicae (Hemiptera: Aphididae). To do so, we used 15 monochromatic lights (emitting various specific wavelengths from 340 to 649 nm) and white light. P. aphidis adults are diurnal insects that show a positive phototaxis to a broad spectrum of light. Significant differences were found between sexes in the phototactic responses of P. aphidis to different monochromatic lights. Female P. aphidis showed four peaks of sensitivity at 380, 450, 504 and 589 nm. Male P. aphidis show two peaks of sensitivity, one at 450 nm and second at 628 nm. P. aphidis adults showed an increased phototactic response at low intensities and a decreased phototactic response at high intensities for both UV light and blue light. This experiment will help provide a scientific basis for the development of colour traps for insect pest management.     

The role of the ocelli in the phototactic behaviour of the haematophagous bug Triatoma infestans

In addition to compound eyes, most adult insects posses two or three simple eyes, the ocelli. The function of these photoreceptors remains elusive in most cases. Triatomine bugs posses two well-developed ocelli, located in a latero-dorsal position, behind the compound eyes. We tested the role of the ocelli in the phototactic behaviour of Triatoma infestans, by measuring the time spent by adult males in the dark half of an experimental arena, which had the other half illuminated. The occlusion of the ocelli or the compound eyes alone had little effect on the phototactic response of the bugs. Only those insects which had both their ocelli and compound eyes occluded showed a significant reduction in their negative response to light. The ability of the ocelli of T. infestans to mediate the phototactic response by themselves (i.e., not through the modulation of compound eyes sensitivity) constitutes the first report on this function in insects.     

光谱和光强度对棕榈蓟马雌成虫行为反应的影响

室内研究了光谱、光强度对棕榈蓟马雌成虫的趋、避光行为的影响。结果显示:在340—605 nm波谱内棕榈蓟马雌成虫对14个单色光刺激的趋光行为反应为多峰型。其中蓝光483 nm处峰最高,趋光反应率达34.96%,其次为绿光498—524 nm、562—582 nm、紫外光340 nm处;其避光行为反应共有3个峰,其中紫外光380 nm处最高,避光率18.08%,另外2个峰分别在橙光605 nm、紫光420 nm处。在趋光率较高的单色光(340、483、524、582 nm)和避光率较高的单色光(380、605 nm)以及白光刺激下,棕榈蓟马雌成虫的趋光率随光强增强的增强而提高,而避光率则随着光强的增强而降低;光强最弱时仍均有一定趋光率,最强时均未出现高端平台。因此:棕榈蓟马雌成虫对不同单色光具有明显的选择性,光谱和光强度对其趋光行为和避光行为有较大影响,光强度的影响作用与波长因素有关。     

信息化合物对昆虫行为的影响

本文综述了来自寄主植物的挥发性物质和同种昆虫或异种昆虫释放的各种信息素及两者的协同作用的信息化合物对昆虫行为的影响。特别强调了寄主植物的气味物质和昆虫信息素协同作用在昆虫寻找寄主、求偶、交配及天敌在寄主识别过程中的重要地位。昆虫对寄主植物的识别是由于识别了植物气味的由一定组分、按照严格比例组成的化学指纹图。昆虫信息素与植物挥发性物质相结合为昆虫寻找求偶、交配场所提供更复杂或更全面的信息。许多昆虫只有在寄主植物或寄主植物气味存在时 ,才能释放性或聚集信息素。天敌在寄主识别、搜索及定位等一系列过程中 ,来自寄主的食料、寄主本身及两者的互作的信息化合物起重要的作用。研究信息化合物对昆虫行为的影响可以探索昆虫各种行为的内在机理 ,更好的了解寄主—昆虫—天敌三层营养关系的相互作用 ,对利用天然活性化合物防治害虫及生物防治提供理论依据     

Phototactic Responses of Cell Population to Repeated Pulses of Yellow Light in a Phytoflagellate Cryptomonas sp

Positive phototaxis in cell populations of a phytoflagellate Cryptomonas sp. was recorded photoelectrically when the duration and intensity of repeated pulses of monochromatic yellow light (570 nm) interspersed with darkness were varied. Irrespective of the duration of the light pulses, phototactic responses to repeated pulses were as great as those to continuous irradiation and were linearly dependent on the logarithm of total incident light energy when the dark interval was shorter than 60 milliseconds. Under these conditions, reciprocity between duration and intensity held well. In contrast, when the dark interval exceeded 250 milliseconds, the responses were remarkably reduced regardless of light duration and were not affected by increasing the intensity of actinic light pulses.

The present results clearly indicate that continuous stimulation with actinic light is not essential for the maximum effect, but that the length of dark interval is crucial in phototactic response.

     

The sensitivity of chlamydomonas photoreceptor is optimized for the frequency of cell body rotation.

For phototactic migration, Chlamydomonas scans the surrounding light environment by rotating the cell body with an eyespot located on the equator. The intensity of the light signal received by the eyespot should therefore change cyclically at the frequency of the cell body rotation. In this study, the response of the photoreceptor to cyclically changing light stimuli was analyzed using immotile mutant cells. To simulate the light intensity change perceived by a rotating cell, light stimuli were applied that consisted of a light phase with the intensity changing similar to a half cycle of a sine wave and a dark phase of the same length. The fluence rate at the peak of the sine wave was of the order of 10(19) photons m(-2) s(-1), i.e. high intensity at which phototaxis is saturated. A photoreceptor current (PRC) was produced at the onset of each light phase. Interestingly, its amplitude varied depending on the frequency and was largest at 1-5 Hz, a frequency range similar to the frequency of cell body rotation. Experiments on the kinetics of the PRC indicate that the response was small at low frequency because of the inactivation of the PRC before full activation. In contrast, at high frequency the PRC was suppressed by adaptation to the repetitive stimuli. These characteristic kinetics of the PRC should be important for Chlamydomonas cells to extract information from the signals generated by the cell body rotation.     

Circadian control of photonegative sensitivity in the haematophagous bug Triatoma infestans

The negative phototactic response of Triatoma infestans (Hemiptera) was studied in an arena, half of which was kept dark and the other half illuminated with different light intensitites. For each intensity we measured the time the insects spent in the dark half, the time to reach the side opposite to that where they were released, and the number of passages through the middle line of the arena. T. infestans displayed a photonegative behaviour that was enhanced by high light intensities. Bugs maintained in 12:12 light-dark cycles responded differently to the same illumination levels when tested in their photophase and scotophase: sensitivity to light was higher during the latter. Bugs entrained to light-dark cycles, kept afterwards either in constant darkness or in constant light, and tested in their subjective night and day, showed the same responses as bugs from the light-dark group tested in their corresponding photophase and scotophase. Thus, phototactic sensitivity is under endogenous control. The behaviour shown by T. infestans may be understood as being composed of at least two different drives: an exploratory one, and a negative phototactic response that is under endogenous control and is particularly sensitive to light during the scotophase, when activity peaks occur. Accepted: 16 July 1998     

Pheromones enhance somatosensory processing in newt brains through a vasotocin-dependent mechanism

We tested whether the sex pheromones that stimulate courtship clasping in male roughskin newts do so, at least in part, by amplifying the somatosensory signals that directly trigger the motor pattern associated with clasping and, if so, whether that amplification is dependent on endogenous vasotocin (VT). Female olfactory stimuli increased the number of action potentials recorded in the medulla of males in response to tactile stimulation of the cloaca, which triggers the clasp motor reflex, as well as to tactile stimulation of the snout and hindlimb. That enhancement was blocked by exposing the medulla to a V1a receptor antagonist before pheromone exposure. However, the antagonist did not affect medullary responses to tactile stimuli in the absence of pheromone exposure, suggesting that pheromones amplify somatosensory signals by inducing endogenous VT release. The ability of VT to couple sensory systems together in response to social stimulation could allow this peptide to induce variable behavioural outcomes, depending on the immediate context of the social interaction and thus on the nature of the associated stimuli that are amplified. If widespread in vertebrates, this mechanism could account for some of the behavioural variability associated with this and related peptides both within and across species.     

A single sex pheromone receptor determines chemical response specificity of sexual behavior in the silkmoth Bombyx mori

In insects and other animals, intraspecific communication between individuals of the opposite sex is mediated in part by chemical signals called sex pheromones. In most moth species, male moths rely heavily on species-specific sex pheromones emitted by female moths to identify and orient towards an appropriate mating partner among a large number of sympatric insect species. The silkmoth, Bombyx mori, utilizes the simplest possible pheromone system, in which a single pheromone component, (E, Z)-10,12-hexadecadienol (bombykol), is sufficient to elicit full sexual behavior. We have previously shown that the sex pheromone receptor BmOR1 mediates specific detection of bombykol in the antennae of male silkmoths. However, it is unclear whether the sex pheromone receptor is the minimally sufficient determination factor that triggers initiation of orientation behavior towards a potential mate. Using transgenic silkmoths expressing the sex pheromone receptor PxOR1 of the diamondback moth Plutella xylostella in BmOR1-expressing neurons, we show that the selectivity of the sex pheromone receptor determines the chemical response specificity of sexual behavior in the silkmoth. Bombykol receptor neurons expressing PxOR1 responded to its specific ligand, (Z)-11-hexadecenal (Z11-16:Ald), in a dose-dependent manner. Male moths expressing PxOR1 exhibited typical pheromone orientation behavior and copulation attempts in response to Z11-16:Ald and to females of P. xylostella. Transformation of the bombykol receptor neurons had no effect on their projections in the antennal lobe. These results indicate that activation of bombykol receptor neurons alone is sufficient to trigger full sexual behavior. Thus, a single gene defines behavioral selectivity in sex pheromone communication in the silkmoth. Our findings show that a single molecular determinant can not only function as a modulator of behavior but also as an all-or-nothing initiator of a complex species-specific behavioral sequence.     

昆虫趋光性及其机理的研究进展

依据目前已有资料 ,从行为学、生理学及田间应用调查等方面概述了近几十年昆虫趋光性的国内外研究进展。其中 ,行为学与生理学研究较多 ,且两者结果基本一致 ,相互补充 ,为趋光性机制的假说提供了可靠的依据。关于昆虫趋光性机制的假说较多 ,其中报道较多的是光干扰假说、光定向行为假说和生物天线假说 3种 ,现在较为普遍接受的是前两者。光干扰假说是指刺眼作用干扰昆虫的正常活动导致趋光 ,而光定向行为假说则指昆虫趋光是由于光定向行为所致     

昆虫定向机制研究进展

许多昆虫具有定向运动的行为。对部分社会性昆虫和迁飞性昆虫定向行为的大量研究已经初步阐明太阳、地磁场、天体、风及地面标志物等都可能成为昆虫返巢和迁飞定向的线索。社会性昆虫具有对不同定向线索进行整合而实现精确导航的能力。日间迁飞性昆虫利用时间补偿太阳罗盘进行定向的机制亦已明确,但夜间迁飞昆虫的定向机制尚需深入研究。迁飞性害虫定向机制的明确将有助于判断害虫迁飞路径及降落区域,为迁飞害虫的准确预测提供科学依据。本文对昆虫的定向机制研究进展进行了综述。     

Probing real sensory worlds of receivers with unsupervised clustering

The task of an organism to extract information about the external environment from sensory signals is based entirely on the analysis of ongoing afferent spike activity provided by the sense organs. We investigate the processing of auditory stimuli by an acoustic interneuron of insects. In contrast to most previous work we do this by using stimuli and neurophysiological recordings directly in the nocturnal tropical rainforest, where the insect communicates. Different from typical recordings in sound proof laboratories, strong environmental noise from multiple sound sources interferes with the perception of acoustic signals in these realistic scenarios. We apply a recently developed unsupervised machine learning algorithm based on probabilistic inference to find frequently occurring firing patterns in the response of the acoustic interneuron. We can thus ask how much information the central nervous system of the receiver can extract from bursts without ever being told which type and which variants of bursts are characteristic for particular stimuli. Our results show that the reliability of burst coding in the time domain is so high that identical stimuli lead to extremely similar spike pattern responses, even for different preparations on different dates, and even if one of the preparations is recorded outdoors and the other one in the sound proof lab. Simultaneous recordings in two preparations exposed to the same acoustic environment reveal that characteristics of burst patterns are largely preserved among individuals of the same species. Our study shows that burst coding can provide a reliable mechanism for acoustic insects to classify and discriminate signals under very noisy real-world conditions. This gives new insights into the neural mechanisms potentially used by bushcrickets to discriminate conspecific songs from sounds of predators in similar carrier frequency bands.     

Circadian mating activity and effect of pheromone pre-exposure on pheromone response rhythms in the moth Spodoptera littoralis

Mating in moths is generally mediated by female-produced sex pheromones. Mating activity, female pheromone production/release and male pheromone responsiveness all show diurnal variations in many species. We found that the response of the male Egyptian cotton leafworm, Spodoptera littoralis, to sex pheromone gland extracts showed a diel rhythm in olfactometer tests, and the variation was persistent for at least 1 day in constant darkness. High male response to sex pheromone was correlated in time with high mating and locomotor activity. Male S. littoralis, maintained in constant darkness and exposed to pheromone gland extracts on a daily basis, showed an induced temporal variation in response after several days, in contrast to unexposed males. This suggests that in the absence of other external zeitgebers, exposure to sex pheromone may function to synchronise circadian behavioural rhythms in male moths. The daily rhythm in mating activity in S. littoralis is also shown to be persistent for at least 2 days in constant darkness. Pairs mated significantly less when either the male or female had been raised in a light:dark cycle 10 h out of phase, indicating that the proposed circadian rhythm in mating activity is composed of rhythmic mating preference/ability in both sexes.     

Phototactic behaviour of the parasitoid Encarsia formosa (Hymenoptera: Aphelinidae)

We investigated the spectral sensitivity and response to light intensity of Encarsia formosa (Hymenoptera: Aphelinidae), which is a key natural enemy of the greenhouse whitefly, Trialeurodes vaporariorum, and the tobacco whitefly, Bemisia tabaci (Hemiptera: Aleyrodidae). To do so, we used 15 monochromatic lights (emitting various specific wavelengths from 340 to 649?nm) and white light. E. formosa adults, which are diurnal insects, showed a positive phototaxis to a broad spectrum of light, with peaks of sensitivity at 414, 340, 450, and 504?nm. These results show that this parasitoid is generally more sensitive to short wavelength lights than long wavelength lights across all spectral ranges tested. Furthermore, E. formosa adults showed an increased phototactic response at low intensities and a decreased response at high intensities, for both ultraviolet light and violet light. Thus, E. formosa showed both colour and intensity preferences. This experiment provides a scientific basis for the development of colour traps for insect pest management and improves understanding of the ecological significance of colour vision by E. formosa.     

昆虫趋光的性别差异及其影响因素

趋光是众多夜行性昆虫的主要行为特征之一.研究表明: 许多夜行性昆虫的趋光存在明显的性别差异现象,而这种差异受多种因素的影响.本文从雌雄成虫飞行能力差异导致其飞行距离与高度的不同、雌雄成虫复眼结构差异导致其对光源反应不同,以及雌雄成虫对光源和环境刺激的敏感性不同等方面综述了导致昆虫趋光性别差异的因素,并探讨了昆虫趋光性别差异的应用及今后研究方向.     

Phototactic Behavior of <Emphasis Type="Italic">Scleroderma guani</Emphasis> (Hymenoptera: Bethylidae) - Parasitoid of <Emphasis Type="Italic">Pissodes punctatus</Emphasis> (Coleoptera: Curculionidae)

Although light trap can be used to control pest populations, they can also kill the natural enemies of pests. Scleroderma guani (Hymenoptera: Bethylidae) is a parasitoid of a bark-weevil Pissodes punctatus (Coleoptera: Curculionidae). To understand the phototactic behavior of S. guani, we investigated its diurnal and nocturnal behavior, then examined its phototactic response to nine monochromatic lights and to five intensities of the two most attractive lights. Our results showed that S. guani is most active during the day, while remain still in a dark room or at night. S. guani showed a positive response both to a broad spectrum of monochromatic light and total light (natural light), which implies a broad sensitivity to the light spectrum. S. guani was most sensitive to blue (450 nm) and green (549 nm) lights, suggesting its visual system composed of blue and green receptors. S. guani was least sensitive to ultraviolet (340 nm) light, which may be caused by long-term mass rearing and propagating under artificial conditions. Furthermore, low intensities elicited a positive phototactic response, while high intensities showed a decreased trend under both blue and green lights. Thus, S. guani is a phototactic insect which shows preferences for light in both color and intensity. This study suggests that light trap can only be utilized to control the adult P. punctatus during and after its peak emergence, due to the overlap in the spectral sensitivity of both pest and parasitoid adults.