The circadian system of the cockroach Leucophaea maderae: The role of the neuropeptide orcokinin and light entrainment

Das circadiane system der Schabe Leucophaea maderae: Die Rolle des Neuropetides Orcokinin und die Synchronisation der Inneren Uhr durch Licht

Processing of sky compass cues and wide-field motion in the central complex of the desert locust (Schistocerca gregaria)

1. Polarization-sensitive neurons of the locust central complex show azimuthdependent responses to unpolarized light spots. This suggests that direct sunlight supports the sky polarization compass in this brain area. / 2. In the brain of the desert locust, neurons sensitive to the plane of celestial polarization are arranged like a compass in the slices of the central complex. These neurons, in addition, code for the horizontal direction of an unpolarized light cue possibly representing the sun. We show here that horizontal directions are, in addition to E-vector orientations from dorsal direction, represented in a compass-like manner across the slices of the central complex. However, both compasses are not linked to each other but seem to interact in a cell specific nonlinear way. Our study confirms the role of the central complex in signaling heading directions signaling and shows that different cues are employed for this task. / 3. Visual cues are essential for animal navigation and spatial orientation. Many insects rely on celestial cues for spatial orientation, including the sky polarization pattern. In desert locusts neurons encoding the plane of polarized light (E-vector) are located in the central complex (CX), a group of midline-spanning neuropils. Several types of CX neuron signalling heading direction represent zenithal Evectors in a topographic manner across the slices of the CX and, likely, act as an internal sky compass. Because animals experience optic flow stimulation during flight, we asked whether progressive wide-field motion affects the responses of CX neurons to polarized light. In most neurons, progressive motion disadapted the response to the preferred E-vector (i.e. the E-vector eliciting strongest firing), whereas the response to the anti-preferred E-vector remained comparatively unaffected. This suggests context-dependent gain modulation in sky compass signalling. Three types of compass neuron were responsive to motion simulating body rotation around the yaw axis. Depending on arborization domains in the CX and rotation direction these neurons were strongly excited or inhibited. As proposed for Drosophila, they may be involved in shifting compass signal activity across the slices of the CX as the animal turns enabling it to keep track of its heading

Spectral properties of identified polarized-light sensitive interneurons in the brain of the desert locust Schistocerca gregaria

Many migrating animals employ a celestial compass mechanism for spatial navigation. Behavioral experimentsin bees and ants have shown that sun compass navigation may rely on the spectral gradient in the sky as well as onthe pattern of sky polarization. While polarized-light sensitive interneurons (POL neurons) have been identifiedin the brain of several insect species, there are at present no data on the neural basis of coding the spectral gradientof the sky. In the present study we have analyzed the chromatic properties of two identified POL neurons in thebrain of the desert locust. Both neurons, termed TuTu1 and LoTu1, arborize in the anterior optic tubercle andrespond to unpolarized light as well as to polarized light. We show here that the polarized-light response of both types of neuron relies on blue-sensitive photoreceptors. Responses to unpolarized light depended on stimulus position and wavelength. Dorsal unpolarized blue light inhibited the neurons, while stimulation from the ipsilateral side resulted in opponent responses to UV light and green light. While LoTu1 was inhibited by UV light and was excited by green light, one subtype of TuTu1 was excited by UV and inhibited by green light. In LoTu1 the sensitivity to polarized light was at least 2 log units higher than the response to unpolarized light stimuli. Taken together, the spatial and chromatic properties of the neurons may be suited to signal azimuthal directions based on a combination of the spectral gradient and thepolarization pattern of the sky

Immunocytochemical localization of amines and GABA in the optic lobe of the Butterfly, Papilio xuthus

Butterflies have sophisticated color vision. While the spectral organization of the compound eye has been well characterized in the Japanese yellow swallowtail butterfly, Papilio xuthus, neural mechanisms underlying its color vision are largely unexplored. Towards a better understanding of signal processing in the visual system of P. xuthus, we used immunocytochemical techniques to analyze the distribution of transmitter candidates, namely, histamine, serotonin, tyramine and γ-aminobutyric acid (GABA). Photoreceptor terminals in the lamina and medulla exhibited histamine immunoreactivity as demonstrated in other insects. The anti-histamine antiserum also labeled a few large medulla neurons. Medulla intrinsic neurons and centrifugal neurons projecting to the lamina showed serotonin immunoreactivity. Tyramine immunostaining was detected in a subset of large monopolar cells (LMCs) in the lamina, transmedullary neurons projecting to the lobula plate, and cell bodies surrounding the first optic chiasma. An anti-GABA antiserum labeled a subset of LMCs and populations of columnar and tangential neurons surrounding the medulla. Each of the four antisera also labeled a few centrifugal neurons that innervate the lobula complex from the central brain, suggesting that they have neuromodulatory roles. A distinctive feature we found in this study is the possibility that tyramine and GABA act as transmitters in LMCs of P. xuthus, which has not been reported in any other insects so far

The Locust Standard Brain: A 3D Standard of the Central Complex as a Platform for Neural Network Analysis

Many insects use the pattern of polarized light in the sky for spatial orientation and navigation. We have investigated the polarization vision system in the desert locust. To create a common platform for anatomical studies on polarization vision pathways, Kurylas et al. (2008) have generated a three-dimensional (3D) standard brain from confocal microscopy image stacks of 10 male brains, using two different standardization methods, the Iterative Shape Averaging (ISA) procedure and the Virtual Insect Brain (VIB) protocol. Comparison of both standardization methods showed that the VIB standard is ideal for comparative volume analysis of neuropils, whereas the ISA standard is the method of choice to analyze the morphology and connectivity of neurons. The central complex is a key processing stage for polarization information in the locust brain. To investigate neuronal connections between diverse central-complex neurons, we generated a higher-resolution standard atlas of the central complex and surrounding areas, using the ISA method based on brain sections from 20 individual central complexes. To explore the usefulness of this atlas, two central-complex neurons, a polarization-sensitive columnar neuron (type CPU1a) and a tangential neuron that is activated during flight, the giant fan-shaped (GFS) neuron, were reconstructed 3D from brain sections. To examine whether the GFS neuron is a candidate to contribute to synaptic input to the CPU1a neuron, we registered both neurons into the standardized central complex. Visualization of both neurons revealed a potential connection of the CPU1a and GFS neurons in layer II of the upper division of the central body

Opsin expression, physiological characterization and identification of photoreceptor cells in the dorsal rim area and main retina of the desert locust, Schistocerca gregaria.

For compass orientation many insects rely on the pattern of sky polarization, but some species also exploit the sky chromatic contrast. Desert locusts, Schistocerca gregaria, detect polarized light through a specialized dorsal rim area (DRA) in their compound eye. To better understand retinal mechanisms underlying visual navigation, we compared opsin expression, spectral and polarization sensitivities and response-stimulus intensity functions in the DRA and main retina of the locust. In addition to previously characterized opsins of long-wavelength-absorbing (Lo1) and blue-absorbing visual pigments (Lo2), we identified an opsin of an ultraviolet-absorbing visual pigment (LoUV). DRA photoreceptors exclusively expressed Lo2, had peak spectral sensitivities at 441 nm and showed high polarization sensitivity (PS 1.3-31.7). In contrast, ommatidia in the main eye co-expressed Lo1 and Lo2 in five photoreceptors, expressed Lo1 in two proximal photoreceptors, and Lo2 or LoUV in one distal photoreceptor. Correspondingly, we found broadband blue- and green-peaking spectral sensitivities in the main eye and one narrowly tuned UV peaking receptor. Polarization sensitivity in the main retina was low (PS 1.3-3.8). V-log I functions in the DRA were steeper than in the main retina, supporting a role in polarization vision. Desert locusts occur as two morphs, a day-active gregarious and a night-active solitarious form. In solitarious locusts, sensitivities in the main retina were generally shifted to longer wavelengths, particularly in ventral eye regions, supporting a nocturnal lifestyle at low light levels. The data support the role of the DRA in polarization vision and suggest trichromatic colour vision in the desert locust

A Distinct Layer of the Medulla Integrates Sky Compass Signals in the Brain of an Insect

Mass migration of desert locusts is a common phenomenon in North Africa and the Middle East but how these insects navigate is still poorly understood. Laboratory studies suggest that locusts are able to exploit the sky polarization pattern as a navigational cue. Like other insects locusts detect polarized light through a specialized dorsal rim area (DRA) of the eye. Polarization signals are transmitted through the optic lobe to the anterior optic tubercle (AOTu) and, finally, to the central complex in the brain. Whereas neurons of the AOTu integrate sky polarization and chromatic cues in a daytime dependent manner, the central complex holds a topographic representation of azimuthal directions suggesting a role as an internal sky compass. To understand further the integration of sky compass cues we studied polarization-sensitive (POL) neurons in the medulla that may be intercalated between DRA photoreceptors and AOTu neurons. Five types of POL-neuron were characterized and four of these in multiple recordings. All neurons had wide arborizations in medulla layer 4 and most, additionally, in the dorsal rim area of the medulla and in the accessory medulla, the presumed circadian clock. The neurons showed type-specific orientational tuning to zenithal polarized light and azimuth tuning to unpolarized green and UV light spots. In contrast to neurons of the AOTu, we found no evidence for color opponency and daytime dependent adjustment of sky compass signals. Therefore, medulla layer 4 is a distinct stage in the integration of sky compass signals that precedes the time-compensated integration of celestial cues in the AOTu

A unified platform to manage, share, and archive morphological and functional data in insect neuroscience

Insect neuroscience generates vast amounts of highly diverse data, of which only a small fraction are findable, accessible and reusable. To promote an open data culture, we have therefore developed the InsectBrainDatabase (IBdb), a free online platform for insect neuroanatomical and functional data. The IBdb facilitates biological insight by enabling effective cross-species comparisons, by linking neural structure with function, and by serving as general information hub for insect neuroscience. The IBdb allows users to not only effectively locate and visualize data, but to make them widely available for easy, automated reuse via an application programming interface. A unique private mode of the database expands the IBdb functionality beyond public data deposition, additionally providing the means for managing, visualizing, and sharing of unpublished data. This dual function creates an incentive for data contribution early in data management workflows and eliminates the additional effort normally associated with publicly depositing research data