Despite the importance of the insect nervous system for functional and developmental neuroscience, descriptions of insect brains have suffered from a lack of uniform nomenclature. Ambiguous definitions of brain regions and fiber bundles have contributed to the variation of names used to describe the same structure. The lack of clearly determined neuropil boundaries has made it difficult to document precise locations of neuronal projections for connectomics study. To address such issues, a consortium of neurobiologists studying arthropod brains, the Insect Brain Name Working Group, has established the present hierarchical nomenclature system, using the brain of Drosophila melanogaster as the reference framework, while taking the brains of other taxa into careful consideration for maximum consistency and expandability. The following summarizes the consortium's nomenclature system and highlights examples of existing ambiguities and remedies for them. This nomenclature is intended to serve as a standard of reference for the study of the brain of Drosophila and other insects.
In the moth Manduca sexta, the number and morphology of neuronal connections between the antennal lobes and the protocerebrum were examined. Cobalt injections revealed eight morphological types of neurons with somata adjacent to the AL neuropil that project in the inner, middle, and outer antenno-cerebral tracts to the proto-cerebrum. Neurons innervating the macroglomerular complex and many neurons with fibers in the inner antenno-cerebral tract have uniglomerular antennal-lobe arborizations. Most neurons in the middle and outer antenno-cerebral tracts, on the other hand, seem to innervate more than one glomerulus. Protocerebral areas receiving direct input from the antennal lobe include the calyces of the mushroom bodies, and circumscribed areas termed "olfactory foci" in the lateral horn of the protocerebrum and several other regions, especially areas in close proximity to the mushroom bodies. Fibers in the inner antenno-cerebral tract that innervate the male-specific macroglomerular complex have arborizations in the protocerebrum that are distinct from the projections of sexually non-specific neurons. Protocerebral neurons projecting into the antennal lobe are much less numerous than antennal-lobe output cells. Most of these protocerebral fibers enter the antennal lobe in small fiber tracts that are different from those described above. In the protocerebrum, these centrifugal cells arborize in olfactory foci and also in the inferior median protocerebrum and the lateral accessory lobes. The morphological diversity of connections between the antennal lobes and the protocerebrum, described here for the first time on a single-cell level, suggests a much greater physiological complexity of the olfactory system than has been assumed so far.
The central complex is a group of neuropils in the center of the insect brain. It consists of four major subunits: the upper and lower divisions of the central body (CBU, CBL), the protocerebral bridge (PB), and the paired noduli. A distinctive feature of the central complex is a modular architecture characterized by rows of 16 columns, intersected in the central body by stacks of layers. Evidence from locusts suggests that the central complex plays a major role in sky compass orientation. To understand signal processing in this brain area further, we have analyzed the morphologies of columnar neurons of the central complex of the locust Schistocerca gregaria. Intracellular dye fills revealed 21 types of columnar neurons that connect columns of different subunits, three types of pontine neurons linking pairs of columns within the CBU, and one amacrine cell. Most neurons appeared to be part of isomorphic sets with cell type-specific heterolateral projection patterns. Evaluation of arborization areas and neuron polarity suggests that these neurons are either intrinsic to the central complex or provide output to the lateral accessory lobes (LALs) or anterior lip region. No direct connections were found between the CBU and CBL. Instead, neurons of either subdivision were connected with the PB, but projected to non-overlapping regions in the LALs and to different layers of the noduli. This study provides novel insights into the functional organization of the central complex, especially with respect to its likely role in right-left signal matching and decision making.
The central complex is a group of modular neuropils across the midline of the insect brain. Hallmarks of its anatomical organization are discrete layers, an organization into arrays of 16 slices along the right-left axis, and precise inter-hemispheric connections via chiasmata. The central complex is connected most prominently with the adjacent lateral complex and the superior protocerebrum. Its developmental appearance corresponds with the appearance of compound eyes and walking legs. Distinct dopaminergic neurons control various forms of arousal. Electrophysiological studies provide evidence for roles in polarized light vision, sky compass orientation, and integration of spatial information for locomotor control. Behavioral studies on mutant and transgenic flies indicate roles in spatial representation of visual cues, spatial visual memory, directional control of walking and flight, and place learning. The data suggest that spatial azimuthal directions (i.e., where) are represented in the slices, and cue information (i.e., what) are represented in different layers of the central complex.
For many insects, the polarization pattern of the blue sky serves as a compass cue for spatial navigation. E-vector orientations are detected by photoreceptors in a dorsal rim area of the eye. Polarized-light signals from both eyes are finally integrated in the central complex, a brain area consisting of two subunits, the protocerebral bridge and the central body. Here we show that a topographic representation of zenithal E-vector orientations underlies the columnar organization of the protocerebral bridge in a locust. The maplike arrangement is highly suited to signal head orientation under the open sky.
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