The Insects has been the standard textbook in the field since the first edition published over forty years ago. Building on the strengths of Chapman's original text, this long-awaited 5th edition has been revised and expanded by a team of eminent insect physiologists, bringing it fully up-to-date for the molecular era. The chapters retain the successful structure of the earlier editions, focusing on particular functional systems rather than taxonomic groups and making it easy for students to delve into topics without extensive knowledge of taxonomy. The focus is on form and function, bringing together basic anatomy and physiology and examining how these relate to behaviour. This, combined with nearly 600 clear illustrations, provides a comprehensive understanding of how insects work. Now also featuring a richly illustrated prologue by George McGavin, this is an essential text for students, researchers and applied entomologists alike.

The insects: structure and function.

Mosaic analysis with a repressible cell marker for studies of gene function in neuronal morphogenesis.

The administration of leptin to leptin-deficient humans, and the analogous Lepob/Lepob mice, effectively reduces hyperphagia and obesity. But common obesity is associated with elevated leptin, which suggests that obese humans are resistant to this adipocyte hormone. In addition to regulating long-term energy balance, leptin also rapidly affects neuronal activity. Proopiomelanocortin (POMC) and neuropeptide-Y types of neurons in the arcuate nucleus of the hypothalamus are both principal sites of leptin receptor expression and the source of potent neuropeptide modulators, melanocortins and neuropeptide Y, which exert opposing effects on feeding and metabolism. These neurons are therefore ideal for characterizing leptin action and the mechanism of leptin resistance; however, their diffuse distribution makes them difficult to study. Here we report electrophysiological recordings on POMC neurons, which we identified by targeted expression of green fluorescent protein in transgenic mice. Leptin increases the frequency of action potentials in the anorexigenic POMC neurons by two mechanisms: depolarization through a nonspecific cation channel; and reduced inhibition by local orexigenic neuropeptide-Y/GABA (gamma-aminobutyric acid) neurons. Furthermore, we show that melanocortin peptides have an autoinhibitory effect on this circuit. On the basis of our results, we propose an integrated model of leptin action and neuronal architecture in the arcuate nucleus of the hypothalamus.

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Leptin activates anorexigenic POMC neurons through a neural network in the arcuate nucleus

Recent advances in integrative studies of locomotion have revealed several general principles. Energy storage and exchange mechanisms discovered in walking and running bipeds apply to multilegged locomotion and even to flying and swimming. Nonpropulsive lateral forces can be sizable, but they may benefit stability, maneuverability, or other criteria that become apparent in natural environments. Locomotor control systems combine rapid mechanical preflexes with multimodal sensory feedback and feedforward commands. Muscles have a surprising variety of functions in locomotion, serving as motors, brakes, springs, and struts. Integrative approaches reveal not only how each component within a locomotor system operates but how they function as a collective whole.

https://science.sciencemag.org/content/sci/288/5463/100.full.pdf

How Animals Move: An Integrative View

We have developed a method for temporal and regional gene expression targeting (TARGET) in Drosophila and show the simultaneous spatial and temporal rescue of a memory defect. The transient expression of the rutabaga-encoded adenylyl cyclase in the mushroom bodies of the adult brain was necessary and sufficient to rescue the rutabaga memory deficit, which rules out a developmental brain defect in the etiology of this deficit and demonstrates an acute role for rutabaga in memory formation in these neurons. The TARGET system offers general utility in simultaneously addressing issues of when and where gene products are required.

https://science.sciencemag.org/content/sci/302/5651/1765.full.pdf

Spatiotemporal rescue of memory dysfunction in Drosophila

The neural circuitry of motion processing in insects, as in primates, involves the segregation of different types of visual information into parallel retinotopic pathways that subsequently are reunited at higher levels. In insects, achromatic, motion-sensitive pathways to the lobula plate are separated from color-processing pathways to the lobula. Further parallel subdivisions of the retinotopic pathways to the lobula plate have been suggested from anatomical observations. Here, we provide direct physiological evidence that the two most prominent of these latter pathways are, indeed, functionally distinct: recordings from the retinotopic pathway defined by small-field bushy T-cells (T4) demonstrate only weak directional selectivity to motion, in striking contrast with previously demonstrated strong directional selectivity in the second, T5-cell, pathway. Additional intracellular recordings and anatomical descriptions have been obtained from other identified neurons that may be crucial in early motion detection and processing: a deep medulla amacrine cell that seems well suited to provide the lateral interactions among retinotopic elements required for motion detection; a unique class of Y-cells that provide small-field, directionally selective feedback from the lobula plate to the medulla; and a new heterolateral lobula plate tangential cell that collates directional, motion-sensitive inputs. These results add important new elements to the set of identified neurons that process motion information. The results suggest specific hypotheses regarding the neuronal substrates for motion-processing circuitry and corroborate behavioral studies in bees that predict distinct pathways for directional and nondirectional motion.

https://www.jneurosci.org/content/jneuro/16/15/4551.full.pdf

Visual Motion-Detection Circuits in Flies: Parallel Direction- and Non-Direction-Sensitive Pathways between the Medulla and Lobula Plate

The shared organization of three optic lobe neuropils—the lamina, medulla, and lobula— linked by chiasmata has been used to support arguments that insects and malacostracans are sister groups. However, in certain insects, the lobula is accompanied by a tectum-like fourth neuropil, the lobula plate, characterized by wide-field tangential neurons and linked to the medulla by uncrossed axons. The identification of a lobula plate in an isopod crustacean raises the question of whether the lobula plate of insects and isopods evolved convergently or are derived from a common ancestor. This question is here investigated by comparisons of insect and crustacean optic lobes. The basal branchiopod crustacean Triops has only two visual neuropils and no optic chiasma. This finding contrasts with the phyllocarid Nebalia pugettensis, a basal malacostracan whose lamina is linked by a chiasma to a medulla that is linked by a second chiasma to a retinotopic outswelling of the lateral protocerebrum, called the protolobula. In Nebalia, uncrossed axons from the medulla supply a minute fourth optic neuropil. Eumalacostracan crustaceans also possess two deep neuropils, one receiving crossed axons, the other uncrossed axons. However, in primitive insects, there is no separate fourth optic neuropil. Malacostracans and insects also differ in that the insect medulla comprises two nested neuropils separated by a layer of axons, called the Cuccati bundle. Comparisons suggest that neuroarchitectures of the lamina and medulla distal to the Cuccati bundle are equivalent to the eumalacostracan lamina and entire medulla. The occurrence of a second optic chiasma and protolobula are suggested to be synapomorphic for a malacostracan/insect clade. J. Comp. Neurol. 467:150 –172, 2003. © 2003 Wiley-Liss, Inc. Indexing terms: evolution; optic lobes; immunocytology; insects; crustaceans

Conserved and convergent organization in the optic lobes of insects and isopods, with reference to other crustacean taxa.

Distribution and neuronal organization of sensilla on the surface of the annulate flagellar segment of the antenna of the maleManduca sexta were studied by scanning and transmission electron microscopy. Nine types of sensilla were identified and their bipolar neurons ascribed to specific sensory modalities on the basis of their cuticular and dendritic morphology. Cuticle morphology identifies two types of sensilla trichodea, two types of sensilla basiconica and one type of sensillum coeloconicum. Certain of these olfactory sensilla are further subdivided on the basis of their dendritic structures. One type of sensillum chaeticum was interpreted as a contact chemoreceptor. A second type of sensillum coeloconicum and a styliform sensilla complex were interpreted as bimodal hygro- and thermosensilla. A second species of sensillum chaeticum serves mechanosensation. Counts from annuli situated about midway along the flagellum revealed a total of about 2200 sensilla supplied by approximately 5160 sensory neurons. A conservative estimate suggests that a male antenna with 85–90 annuli provides the flagellar nerve with at least 3.6 × 105 receptor axons, a number that exceeds previous estimates by almost 50% Each species of receptor has a characteristic location on the annulus. Of the 2100 or so sensilla situated on the dorsal, ventral and the leading edge surfaces, about 800 consist of male-specific type-I trichoids containing pheromone-sensitive receptors. Arciform arrays of these sensilla on the upper and lower surfaces of each annulus presumably optimize the capture and absorbtion of odour molecules. The trailing edge of the flagellum, which is thickly covered by scales and was assumed until now to lack receptors, contains both mechanosensitive and contact chemoreceptors. The modality of non-olfactory receptors is considered with respect to similar elements that have been functionally identified in other species. The coexistence of non-olfactory sensilla with olfactory elements is discussed with respect to current knowledge of the organization of olfactory centres in the brain.

Structure, distribution and number of surface sensilla and their receptor cells on the olfactory appendage of the male moth Manduca sexta.

Giant motion-sensitive tangential neurons in the lobula plate are thought to be cardinal elements in the oculomotor pathways of flies. However, these large neurons do not themselves compute motion, and elementary motion detectors have been proposed only from theory. Here we identify the forms, projections, and responses of small-field retinotopic neurons that comprise a well known pathway from the retina to the lobula plate. Already at the level of the second and third synapses beneath the photoreceptor layer, certain of these small elements show responses that distinguish motion from flicker. At a level equivalent to the vertebrate inner plexiform layer (the fly's outer medulla) at least one retinotopic element is directionally selective. At the inner medulla, small retinotopic neurons with bushy dendrites extending through a few neighboring columns leave the inner medulla and supply inputs onto lobula plate tangentials. These medulla relays have directionally selective responses that are indistinguishable from those of large-field tangentials except for their amplitude and modulation with contrast frequency. Centrifugal neurons leading back from the inner medulla out to the lamina also show orientation-selective responses to motion. The results suggest that specific cell types between the lamina and inner medulla correspond to stages of the Hassenstein-Reichardt correlation model of motion detection.

https://www.jneurosci.org/content/jneuro/15/8/5596.full.pdf

Visual motion detection circuits in flies: peripheral motion computation by identified small-field retinotopic neurons

Certain intact nerve cells in flies can be filled with cobalt from presynaptic or postsynaptic neurons. This cobalt coupling is best demonstrated in giant fibre systems where the phenomenon was originally termed ‘transsynaptic staining’. Fine structural analysis of silver-intensified, cobalt-coupled neurons indicates that the passage of cobalt ions occurs at gap junctions that are accompanied by conventional chemical synapses. Cobalt-coupled systems in dipterous insects are uniquely identifiable and can always be detected between the same kinds of neurons. The visualization of cobalt-coupled neurons allows the identification of functional pathways linking the brain to motor neuropils.

Nicholas J. Strausfeld

Papers

Visual Motion-Detection Circuits in Flies: Parallel Direction- and Non-Direction-Sensitive Pathways between the Medulla and Lobula Plate

Conserved and convergent organization in the optic lobes of insects and isopods, with reference to other crustacean taxa.

Structure, distribution and number of surface sensilla and their receptor cells on the olfactory appendage of the male moth Manduca sexta.

Visual motion detection circuits in flies: peripheral motion computation by identified small-field retinotopic neurons

Cobalt-coupled neurons of a giant fibre system in Diptera