Expression of UV-, blue-, long-wavelength-sensitive opsins and melatonin in extraretinal photoreceptors of the optic lobes of hawkmoths

Lepidopterans display biological rhythms associated with egg laying, eclosion and flight activity but the photoreceptors that mediate these behavioural patterns are largely unknown. To further our progress in identifying candidate light-input channels for the lepidopteran circadian system, we have developed polyclonal antibodies against ultraviolet (UV)-, blue- and extraretinal long-wavelength (LW)-sensitive opsins and examined opsin immunoreactivity in the adult optic lobes of four hawkmoths, Manduca sexta, Acherontia atropos, Agrius convolvuli and Hippotion celerio. Outside the retina, UV and blue opsin protein expression is restricted to the adult stemmata, with no apparent expression elsewhere in the brain. Melatonin, which is known to have a seasonal influence on reproduction and behaviour, is expressed with opsins in adult stemmata together with visual arrestin and chaoptin. By contrast, the LW opsin protein is not expressed in the retina or stemmata but rather exhibits a distinct and widespread distribution in dorsal and ventral neurons of the optic lobes. The lamina, medulla, lobula and lobula plate, accessory medulla and adjacent neurons innervating this structure also exhibit strong LW opsin immunoreactivity. Together with the adult stemmata, these neurons appear to be functional photoreceptors, as visual arrestin, chaoptin and melatonin are also co-expressed with LW opsin. These findings are the first to suggest a role for three spectrally distinct classes of opsin in the extraretinal detection of changes in ambient light and to show melatonin-mediated neuroendocrine output in the entrainment of sphingid moth circadian and/or photoperiodic rhythms.This work was partially supported by the Canadian Institute for Advanced Research (A.D.B.) and the National Science Foundation (grant nos. IBN-0082700 and IBN-0346765; A.D.B.).     

Phylogeny of the Myriapoda – Crustacea – Insecta: a new attempt using photoreceptor structure*

According to molecular sequence data Crustacea and not Myriapoda seem to be the sister‐group to Insecta. This makes it necessary to reconsider how the morphology of their eyes fit with these new cladograms. Homology of facetted eye structures in Insecta (Hexapoda in the sense of Ento‐ and Ectognatha) and Crustacea is clearly supported by identical numbers of cells in an ommatidium (two corneageneous or primary pigment cells, four Semper cells which build the crystalline cone and primarily eight retinula cells). These cell numbers are retained even when great functional modification occurs, especially in the region of the dioptric apparatus. There are two different possibilities to explain differences in eye structure in Myriapoda depending on their phylogenetic position in the cladogram of Mandibulata. In the traditional Tracheata cladogram, eyes of Myriapoda must be secondarily modified. This modification can be explained using the different evolutionary pathways of insect facetted eyes to insect larval eyes (stemmata) as an analogous model system. Comparative morphology of larval insect eyes from all holometabolan orders shows that there are several evolutionary pathways which have led to different types of stemmata and that the process always involved the breaking up the compound eye into individual larval ommatidia. Further evolution led on many occasions to so‐called fusion‐stemmata that occur convergently in each holometabolic order and reveals, in part, great structural similarities to the lateral ocelli of myriapods. As myriapodan eyes cannot be regarded as typical mandibulate ommatidia, their structure can be explained as a modified complex eye evolved in a comparable way to the development to the fusion‐stemmata of insect larvae. The facetted eyes of Scutigera (Myriapoda, Chilopoda) must be considered as secondarily reorganized lateral myriapodan stemmata, the so‐called ‘pseudo‐compound eyes’. New is a crystalline cone‐like vitreous body within the dioptric apparatus. In the new cladogram with Crustacea and Insecta as sister‐groups however, the facetted eyes of Scutigera can be interpreted as an old precursor of the Crustacea – Insecta facetted eye with modified ommatidia having a four‐part crystalline cone, etc. as a synapomorphy. Lateral ocelli of all the other Myriapoda are then modified like insect stemmata. The precursor is then the Scutigera‐Ommatidium. In addition further interpretations of evolutionary pathways of myriapodan morphological characters are discussed.     

Immobile and tough versus mobile and weak: effects of ultraviolet B radiation on eggs and larvae of Manduca sexta

Although indirect effects of solar ultraviolet (UV) radiation on insects are well known (e.g. UV radiation can modify plant chemistry), direct effects of solar radiation on insects have received little attention. Radiation in the UVB range (300–320 nm) is damaging because it is absorbed directly by proteins and DNA. UVB should be toughest on immobile or small life stages, such as eggs or early larval instars. In the present study, the effects of UVB radiation on eggs and larvae of the tobacco hornworm Manduca sexta L. (Lepidoptera: Sphingidae) are examined. The present study aimed to address: what natural levels of UV do they experience; how does UVB affect the performance of eggs; and how does it affect the performance of larvae? In addition, do M. sexta larvae use behaviour to avoid UVB exposure and, consequently, are they physiologically less robust to UVB? In these experiments, eggs and late larval instars of M. sexta are found to be robust to natural levels of UV radiation. By contrast, young larvae are not only more susceptible to damage from UVB, but also they use behavioural means to avoid it. The strategy of using behaviour may relax selection pressures on morphological and physiological mechanisms for preventing (or recovering from) damage by environmental UV radiation.     

Vision in the mantispid: a sit‐and‐wait and stalking predatory insect

Mantispids (Neuroptera: Mantispidae) are remarkable insects as a result of their close resemblance to the praying mantis (order Mantodea). Although not closely related phylogenetically, as a result of similar selective pressures, both mantispids and mantids have evolved powerful raptorial forelegs for capturing insects. Another striking feature is the hypermetamorphosis in mantispid development, as well as the parasitizing behaviour of the first‐instar larvae. The present review focuses on the role of mantispid vision. First, the morphology and functional significance of the larval eyes (stemmata) are examined. In principle, the stemmata are suitable for spatial vision because of their arrangement and structure. This is then followed by a discussion of how adult mantispids are able to capture fast‐moving insects successfully, although, in contrast to the praying mantis, mantispids rely on superposition eyes rather than on apposition eyes with a frontal region of high acuity. For both larvae and adults, comparisons are made with other insect groups. The present review also addresses the role of mantispid vision as an important cue for triggering mating behaviour; accordingly, sex‐specific differences are considered. Finally, vision in the context of orientation flight is discussed.     

昆虫单眼的结构和功能

大多数昆虫的视觉器官除了复眼外还有一些简单的小眼,称为单眼。昆虫成虫和半变态类若虫的单眼称为背单眼,位于头顶两复眼之间。背单眼在数目和结构上都有较大变化,但基本结构包括角膜晶体、一层角膜生成细胞(覆盖在角膜晶体上)、视网膜(由大约1000个感光细胞构成,视类群而不同)。背单眼对弱光比较敏感,但在图像感知方面的作用并不显著;它是一种“激发器官”,可以增加复眼的感知能力。全变态昆虫的幼虫既没有复眼也没有背单眼,但在其头部两侧有些类似复眼小眼的侧单眼。侧单眼的结构也与小眼相似,包括角膜,晶体和由一些视网膜细胞组成的视杆。侧单眼是完全变态类昆虫幼虫仅有的感光器官,与复眼一样,它们可以感知颜色、形状、距离等等。     

Visual behaviour of neonata larvae of the light brown apple moth

When neonate larvae of a leafroller moth,Epiphyas postvittana (Walker) (Lepidoptera: Tortricidae) were released into the middle of a circular arena with blue paper on one side of the arena and apple foliage on the other side, more larvae walked towards the apple foliage. These oriented responses were enhanced, in terms of the number of larvae responding, by increasing the amount of light reflected from or transmitted through apple foliage. Larvae also responded to painted targets, and specifically to targets reflecting light in the region of 470–570 nm (green-yellow region to the human eye). When the amount of 470–570 nm reflected from targets was reduced, numbers of larvae responding to targets decreased. The addition of 400–500 nm reflected light to 470–570 nm reflected light also resulted in a reduction of oriented responses to targets. Tests using neutral colours (white, black, and a series of greys) indicated that, in the absence of targets reflecting primarily in the 470–570 nm region, larvae oriented towards targets with low levels of reflectance. When the orientation of walking larvae was measured at various distances from targets of different colours or diameters, targets subtending 5–8 degrees elicited responses from 50% of all larvae. Behaviour other than walking was also influenced by visual stimuli: fewer larvae spun down on a silken thread when blue paper was placed beneath a walking platform than when brown or green papers were present.     

Visual determinants of escape in tiger beetle larvae (Cicindelidae)

Larvae of tiger beetles (Coleoptera: Cicindelidae) are burrow-dwelling, visual ambush predators which withdraw into their burrows with the passing of large objects. Laboratory experiments confirmed that stimulation of each of the four principal stemmata can elicit escape and that the necessary visual stimulus is contracting, expanding, or transverse movement of a high-contrast image. Response frequency increases as a power function of contrast. Whole-field dimming is ineffective. Movement of large images composed of multiple texture elements, e.g., checkerboards, does not elicit escape, even if each element is much larger than the system's minimum visible angle (4–8° depending upon image contrast). In pilot experiments with a single figure before a textured background, coherent movement of the two inhibits escape, whereas motion in opposite directions does not. Thus, the processing mechanism functions as a feature detector and directs a response to large, single, moving objects.