Optic lobe development.

The optic lobes comprise approximately half of the fly's brain. In four major synaptic ganglia, or neuropils, the visual input from the compound eyes is received and processed for higher order visual functions like motion detection and color vision. A common characteristic of vertebrate and invertebrate visual systems is the point-to-point mapping of the visual world to synaptic layers in the brain, referred to as visuotopy. Vision requires the parallel extraction of numerous parameters in a visuotopic manner. Consequently, the optic neuropils are arranged in columns and perpendicularly oriented synaptic layers that allow for the selective establishment of synapses between columnar neurons. How this exquisite synaptic specificity is established during approximately 100 hours of brain development is still poorly understood. However, the optic lobe contains one of the best characterized brain structures in any organism-both anatomically and developmentally. Moreover, numerous molecules and their function illuminate some of the basic mechanisms involved in brain wiring. The emerging picture is that the development of the visual system of Drosophila is (epi-)genetically hard-wired; it supplies the emerging fly with vision without requiring neuronal activity for fine tuning of neuronal connectivity. Elucidating the genetic and cellular principles by which gene activity directs the assembly of the optic lobe is therefore a fascinating task and the focus of this chapter.

Cylindrical coordinate machining of optical freeform surfaces.

The cylindrical coordinate machining method (CCM) is systematically studied in generating optical freeform surfaces, in which the feature points are fitted to typical Non-Uniform Rational B-Splines (NURBS). The given points have the mapping coordinates in the variable space using the point inversion technique, while the other points have their NURBS coordinates due to the interpolation technique. The derivation and mathematical features are obtained using the fitting formula. The compensation and optimized values for tool geometry are studied using a proposed sectional curve method for fabricating designed surfaces. Typical freeform surfaces fabricated by the CCM method are presented.

Head morphology of Caurinus (Boreidae, Mecoptera) and its phylogenetic implications.

External and internal head structures of Caurinus dectes were examined and described in detail. The features are compared to conditions found in other groups of Antliophora. Caurinus is obviously crucial for the reconstruction of the mecopteran and antliophoran groundplan. It displays a remarkable series of plesiomorphic character states such as a complete clypeolabral suture, the presence of M. hypopharyngomandibularis (M. 13) and M. frontohypopharyngalis (M. 41), a subdivided clypeus, a short head without rostrum, a dorsal tentorial arm attached to the head capsule, the absence of a cranial dilator of the antenna, and large mandibles with a well developed apical tooth, two distinct subapical teeth, and a basal molar part. The first three plesiomorphic features render potential autapomorphies of Mecoptera in the traditional sense invalid. Autapomorphies of Caurinus are the distinctly flattened labrum, the absence of the labroepipharyngeal muscle, the very large size of M. 13, the strongly enlarged penultimate palpomeres, the partition of M. 41, the very strongly developed precerebral sucking chamber, strongly curved optic lobes, the presence of a large protocerebral extension in the genal region and deep posterior excavations of the protocerebrum. The maxillolabial plate, the absence of cardines as separate structures, the reduction of ocelli, and the origin of maxillary palp muscles on a median ridge or area of the maxillolabial plate are likely autapomorphies of Boreidae. Another potential autapomorphy of the family is the presence of longitudinal furrows on the mandibles. However, they are absent in Boreus. The thick strongly sclerotised, median ridge of the maxillolabial plate, the missing retractibility of the prementum, the absence of extrinsic labial muscles, and the presence of a median ridge on the prepharyngeal roof suggest a clade Boreus+Hesperoboreus. The origin of extrinsic maxillary muscles from the clypeus has probably evolved independently in Boreus and Hesperoboreus, and in Panorpa, respectively. The absence of M. craniolacinialis and the presence of a row of several subapical mandibular teeth are autapomorphies of Boreus. The presence of a specific intrinsic muscle of the salivary duct and a membranous galea enclosing the labrum and mandibular base are derived features shared by Boreidae and Pistillifera (galea absent in Nannochorista, Siphonaptera and Diptera). The loss of M. frontolabralis (M. 8) is a potential apomorphy of Mecoptera incl. Siphonaptera. A sister group relationship between Boreidae and Siphonaptera is not supported by characters of the adult head. Head structures of Siphonaptera are extremely modified in correlation with ectoparasitic habits.

Cofilin/ADF is required for retinal elongation and morphogenesis of the Drosophila rhabdomere.

Drosophila photoreceptors undergo marked changes in their morphology during pupal development. These changes include a five-fold elongation of the retinal cell body and the morphogenesis of the rhabdomere, the light sensing structure of the cell. Here we show that twinstar (tsr), which encodes Drosophila cofilin/ADF (actin-depolymerizing factor), is required for both of these processes. In tsr mutants, the retina is shorter than normal, the result of a lack of retinal elongation. In addition, in a strong tsr mutant, the rhabdomere structure is disorganized and the microvilli are short and occasionally unraveled. In an intermediate tsr mutant, the rhabdomeres are not disorganized but have a wider than normal structure. The adherens junctions connecting photoreceptor cells to each other are also found to be wider than normal. We propose, and provide data supporting, that these wide rhabdomeres and adherens junctions are secondary events caused by the inhibition of retinal elongation. These results provide insight into the functions of the actin cytoskeleton during morphogenesis of the Drosophila eye.

Circular polarization vision in a stomatopod crustacean.

We describe the addition of a fourth visual modality in the animal kingdom, the perception of circular polarized light. Animals are sensitive to various characteristics of light, such as intensity, color, and linear polarization [1, 2]. This latter capability can be used for object identification, contrast enhancement, navigation, and communication through polarizing reflections [2-4]. Circularly polarized reflections from a few animal species have also been known for some time [5, 6]. Although optically interesting [7, 8], their signal function or use (if any) was obscure because no visual system was known to detect circularly polarized light. Here, in stomatopod crustaceans, we describe for the first time a visual system capable of detecting and analyzing circularly polarized light. Four lines of evidence-behavior, electrophysiology, optical anatomy, and details of signal design-are presented to describe this new visual function. We suggest that this remarkable ability mediates sexual signaling and mate choice, although other potential functions of circular polarization vision, such as enhanced contrast in turbid environments, are also possible [7, 8]. The ability to differentiate the handedness of circularly polarized light, a visual feat never expected in the animal kingdom, is demonstrated behaviorally here for the first time.

The nauplius eye complex in 'conchostracans'(Crustacea, Branchiopoda: Laevicaudata, Spinicaudata, Cyclestherida) and its phylogenetic implications.

The nauplius eye in Cyclestherida, Laevicaudata and Spinicaudata (previously collectively termed Conchostraca) consists of four cups of inverse sensory cells separated by a pigment layer and a tapetum layer. There are two lateral and two medial cups, a ventral medial cup and a posterior medial cup. The pigment and tapetum layers contain two different kinds of pigment granules, the inner pigment layer relatively large, dark (and electron dense) granules, and the outer tapetum layer light, reflective pigment granules. The presence of four cups and two different kinds of pigment granules are interpreted as autapomorphies of Phyllopoda. The position and shape of the nauplius eye in Spinicaudata is very distinct and herein interpreted as an autapomorphy of this taxon. Additional frontal eyes might be present dorsally or ventrally in varying proximity to the nauplius eye, but they have separate nerves from their sensory cells to the nauplius eye centre in the protocerebrum. Rhabdomeric structures are present in all these frontal eyes, evidencing their light sensitivity. In Lynceus biformis and L. tatei (Laevicaudata), two pairs of frontal eyes were found. In Cyclestheria hislopi (Cyclestherida), an unpaired ventral frontal eye is present. We did not find additional frontal eyes in Limnadopsis parvispinus and Caenestheriella sp. (Spinicaudata).

Phototransduction and retinal degeneration in Drosophila.

Drosophila visual transduction is the fastest known G-protein-coupled signaling cascade and has therefore served as a genetically tractable animal model for characterizing rapid responses to sensory stimulation. Mutations in over 30 genes have been identified, which affect activation, adaptation, or termination of the photoresponse. Based on analyses of these genes, a model for phototransduction has emerged, which involves phosphoinoside signaling and culminates with opening of the TRP and TRPL cation channels. Many of the proteins that function in phototransduction are coupled to the PDZ containing scaffold protein INAD and form a supramolecular signaling complex, the signalplex. Arrestin, TRPL, and G alpha(q) undergo dynamic light-dependent trafficking, and these movements function in long-term adaptation. Other proteins play important roles either in the formation or maturation of rhodopsin, or in regeneration of phosphatidylinositol 4,5-bisphosphate (PIP2), which is required for the photoresponse. Mutation of nearly any gene that functions in the photoresponse results in retinal degeneration. The underlying bases of photoreceptor cell death are diverse and involve mechanisms such as excessive endocytosis of rhodopsin due to stable rhodopsin/arrestin complexes and abnormally low or high levels of Ca2+. Drosophila visual transduction appears to have particular relevance to the cascade in the intrinsically photosensitive retinal ganglion cells in mammals, as the photoresponse in these latter cells appears to operate through a remarkably similar mechanism.

Phototransduction in ganglion-cell photoreceptors.

A third class of photoreceptors has recently been identified in the mammalian retina. They are a rare cell type within the class of ganglion cells, which are the output cells of the retina. These intrinsically photosensitive retinal ganglion cells support a variety of physiological responses to daylight, including synchronization of circadian rhythms, modulation of melatonin release, and regulation of pupil size. The goal of this review is to summarize what is currently known concerning the cellular and biochemical basis of phototransduction in these cells. I summarize the overwhelming evidence that melanopsin serves as the photopigment in these cells and review the emerging evidence that the downstream signaling cascade, including the light-gated channel, might resemble those found in rhabdomeric invertebrate photoreceptors.

Signal integration during development: mechanisms of EGFR and Notch pathway function and cross-talk.

Metazoan development relies on a highly regulated network of interactions between conserved signal transduction pathways to coordinate all aspects of cell fate specification, differentiation, and growth. In this review, we discuss the intricate interplay between the epidermal growth factor receptor (EGFR; Drosophila EGFR/DER) and the Notch signaling pathways as a paradigm for signal integration during development. First, we describe the current state of understanding of the molecular architecture of the EGFR and Notch signaling pathways that has resulted from synergistic studies in vertebrate, invertebrate, and cultured cell model systems. Then, focusing specifically on the Drosophila eye, we discuss how cooperative, sequential, and antagonistic relationships between these pathways mediate the spatially and temporally regulated processes that generate this sensory organ. The common themes underlying the coordination of the EGFR and Notch pathways appear to be broadly conserved and should, therefore, be directly applicable to elucidating mechanisms of information integration and signaling specificity in vertebrate systems.

Odd-skipped genes specify the signaling center that triggers retinogenesis in Drosophila.

Although many of the factors responsible for conferring identity to the eye field in Drosophila have been identified, much less is known about how the expression of the retinal ;trigger', the signaling molecule Hedgehog, is controlled. Here, we show that the co-expression of the conserved odd-skipped family genes at the posterior margin of the eye field is required to activate hedgehog expression and thereby the onset of retinogenesis. The fly Wnt1 homologue wingless represses the odd-skipped genes drm and odd along the anterior margin and, in this manner, spatially restricts the extent of retinal differentiation within the eye field.

Casting a genetic light on the evolution of eyes.

Light has been exploited for information by organisms through the evolution of photoreceptors and, ultimately, eyes in animals. Only a handful of eye types exist because the physics of light constrains photodetection. In the past few years, genetic tools have revealed several parallel pathways through which light guides behavior and have provided insights into the convergent evolution of eyes. The gene encoding opsin (the primary phototransduction protein) and some developmental genes had very early origins and were recruited repeatedly during eye evolution. Eye lens proteins arose separately and make up a diverse group, many of which were co-opted from other functions. A major challenge now is understanding how newly discovered pathways for processing light evolved and how they collaborate with eyes to harvest information from light.

Identification of the Drosophila skpA gene as a novel target of the transcription factor DREF.

SKPa is component of a Drosophila SCF complex that functions in combination with the ubiquitin-conjugating enzyme UbcD1. skpA null mutation results in centrosome overduplication, unusual chromatin condensation, defective endoreduplication and cell-cycle progression. While the molecular mechanisms that regulate expression of the skpA gene are poorly understood, the DNA replication-related element (DRE) and the DRE-binding factor (DREF) play important roles in regulating proliferation-related genes in Drosophila and DRE (5'-TATCGATA) and DRE-like (5'-CATCGATT) sequences were here found to be involved in skpA promoter activity. Thus both luciferase transient expression assays in cultured Drosophila S2 cells using skpA promoter-luciferase fusion plasmids and anti-lacZ immunostaining of various tissues from transgenic third instar larvae carrying the skpA promoter-lacZ fusion genes provided supportive evidence. Furthermore, anti-SKPa immunostaining of eye imaginal discs from flies overexpressing DREF showed ectopic expression of protein in the region posterior to the morphogenetic furrow where DREF is overexpressed. Knockdown of DREF in some tissues where SKPa distribution is well known almost completely abrogated the skpA gene expression. These findings, taken together, indicate that the Drosophila skpA gene is a novel target of the transcription factor DREF.

The Hippo pathway regulates the bantam microRNA to control cell proliferation and apoptosis in Drosophila.

The Hippo signaling pathway acts upon the Yorkie transcriptional activator to control tissue growth in Drosophila. Activated Yorkie drives growth by stimulating cell proliferation and inhibiting apoptosis, but how it achieves this is not understood. Yorkie is known to activate Cyclin E (CycE) and the apoptosis inhibitor DIAP1. However, overexpression of these targets is not sufficient to cause tissue overgrowth. Here we show that Yorkie also activates expression of the bantam microRNA, a known regulator of both proliferation and apoptosis. bantam overexpression mimics Yorkie activation while loss of bantam function slows the rate of cell proliferation. bantam is necessary for Yorkie-induced overproliferation and bantam overexpression is sufficient to rescue survival and proliferation of yorkie mutant cells. Finally, we show that bantam levels are regulated during both developmentally programmed proliferation arrest and apoptosis. In summary, the results show that the Hippo pathway regulates expression of bantam to control tissue growth in Drosophila.

Displacement of D1, HP1 and topoisomerase II from satellite heterochromatin by a specific polyamide.

The functions of DNA satellites of centric heterochromatin are difficult to assess with classical molecular biology tools. Using a chemical approach, we demonstrate that synthetic polyamides that specifically target AT-rich satellite repeats of Drosophila melanogaster can be used to study the function of these sequences. The P9 polyamide, which binds the X-chromosome 1.688 g/cm3 satellite III (SAT III), displaces the D1 protein. This displacement in turn results in a selective loss of HP1 and topoisomerase II from SAT III, while these proteins remain bound to the adjacent rDNA repeats and to other regions not targeted by P9. Conversely, targeting of (AAGAG)n satellite V repeats by the P31 polyamide results in the displacement of HP1 from these sequences, indicating that HP1 interactions with chromatin are sensitive to DNA-binding ligands. P9 fed to larvae suppresses the position-effect variegation phenotype of white-mottled adult flies. We propose that this effect is due to displacement of the heterochromatin proteins D1, HP1 and topoisomerase II from SAT III, hence resulting in stochastic chromatin opening and desilencing of the nearby white gene.

Molecular mechanisms regulating molting in a crustacean.

Crustacean growth and development is characterized by periodic shedding (ecdysis) and replacement of the rigid exoskeleton. Secretions of the X-organ sinus gland complex control the cellular events that lead to growth and molting. Western blot and ELISA results showed a progressive increase in growth arrest-specific protein (Gas7) from early postmolt stage to a maximum at late postmolt stage. Phosphorylation of ERK2, a downstream signaling protein, was also identified in the subsequent stages. ERK2 phosphorylation resulted in the expression of molt-inhibiting hormone (MIH). Specific ERK inhibitors (PD98059 and UO126) exhibited the ability to reduce the molting duration of Fenneropenaeus indicus from 12-14 days to 7-8 days, suggesting that the ERK1/2 signaling pathway is responsible for the expression of MIH, which controls the molt cycle. We have identified the stage-specific expression of Gas7 (approximately 48 kDa) in the X-organ sinus gland complex of eyestalk which is involved in the downstream signaling of the ERK1/2 pathway regulating the expression of MIH during the molt cycle of the white shrimp, F. indicus. These are the first data showing an association between the Gas7 signal-transduction process and regulation of the molt cycle and provides an alternative molecular intervention mechanism to the traditional eyestalk ablation in crustaceans.

Roles of glia in the Drosophila nervous system.

Glial cells have diverse functions that are necessary for the proper development and function of complex nervous systems. Various insects, primarily the fruit fly Drosophila melanogaster and the moth Manduca sexta, have provided useful models of glial function during development. The present review will outline evidence of glial contributions to embryonic, visual, olfactory and wing development. We will also outline evidence for non-developmental functions of insect glia including blood-brain-barrier formation, homeostatic functions and potential contributions to synaptic function. Where relevant, we will also point out similarities between the functions of insect glia and their vertebrate counterparts.

The mechanisms and molecules that connect photoreceptor axons to their targets in Drosophila.

The development of the Drosophila visual system provides a framework for investigating how circuits assemble. A sequence of reciprocal interactions amongst photoreceptors, target neurons and glia creates a precise pattern of connections while reducing the complexity of the targeting process. Both afferent-afferent and afferent-target interactions are required for photoreceptor (R cell) axons to select appropriate synaptic partners. With the identification of some critical cell adhesion and signaling molecules, the logic by which axons make choices amongst alternate synaptic partners is becoming clear. These studies also provide an opportunity to examine the molecular basis of neural circuit evolution.

Physiological characterization of the compound eye in monarch butterflies with focus on the dorsal rim area.

The spectral, angular and polarization sensitivities of photoreceptors in the compound eye of the monarch butterfly (Danaus plexippus) are examined using electrophysiological methods. Intracellular recordings reveal a spectrally homogenous population of UV receptors with optical axes directed upwards and >or=10 degrees to the contralateral side. Based on optical considerations and on the opsin expression pattern (Sauman et al. 2005), we conclude that these UV receptors belong to the anatomically specialized dorsal rim area (DRA) of the eye. Photoreceptors in the main retina with optical axes <10 degrees contralateral or ipsilateral have maximal sensitivities in the UV (lambda(max)

Comparative study on the relationship between photoperiodic host-seeking behavioral patterns and the eye parameters of mosquitoes.

Relationships between the ommatidial structure and photoperiodic behavior of several mosquito species were investigated. Host-seeking behavioral patterns of mosquitoes were classified into four main groups based on previously compiled reports on field or laboratory biting activity. These groups were pattern I and I' (nocturnal), pattern II (crepuscular and nocturnal), pattern III (crepuscular and diurnal), and pattern IV (diurnal). Eye parameters (product of facet diameter and interommatidial angle) of mosquitoes that belong to the pattern I and I' group were higher (2.7-4.2) than those of mosquitoes that belong to the pattern IV group (0.8-2.3). Eye parameters of the mosquitoes categorized in the pattern II and III groups were intermediate (2.3-2.6). These results suggest that the crepuscular behavior of mosquitoes undergoes a transition in the course of evolution from nocturnal behavior to diurnal behavior. Large variations in the eye parameters were observed even within the same genus depending on their photoperiodic behavior. Therefore, the ommatidial structure of mosquitoes appears to be determined, not taxonomically, but evolutionarily by the photoenvironment in which the mosquitoes are most active.