Synaptogenesis in the giant-fibre system of Drosophila: interaction of the giant fibre and its major motorneuronal target

The tergotrochanteral (jump) motorneuron is a major synaptic target of the Giant Fibre in Drosophila. These two neurons are major components of the fly's Giant-Fibre escape system. Our previous work has described the development of the Giant Fibre in early metamorphosis and the involvement of the shaking-B locus in the formation of its electrical synapses. In the present study, we have investigated the development of the tergotrochanteral motorneuron and its electrical synapses by transforming Drosophila with a Gal4 fusion construct containing sequences largely upstream of, but including, the shaking-B(lethal) promoter. This construct drives reporter gene expression in the tergotrochanteral motorneuron and some other neurons. Expression of green fluorescent protein in the motorneuron allows visualization of its cell body and its subsequent intracellular staining with Lucifer Yellow. These preparations provide high-resolution data on motorneuron morphogenesis during the first half of pupal development. Dye-coupling reveals onset of gap-junction formation between the tergotrochanteral motorneuron and other neurons of the Giant-Fibre System. The medial dendrite of the tergotrochanteral motorneuron becomes dye-coupled to the peripheral synapsing interneurons between 28 and 32 hours after puparium formation. Dye-coupling between tergotrochanteral motorneuron and Giant Fibre is first seen at 42 hours after puparium formation. All dye coupling is abolished in a shaking-B(neural) mutant. To investigate any interactions between the Giant Fibre and the tergotroachanteral motorneuron, we arrested the growth of the motorneuron's medial neurite by targeted expression of a constitutively active form of Dcdc42. This results in the Giant Fibre remaining stranded at the midline, unable to make its characteristic bend. We conclude that Giant Fibre morphogenesis normally relies on fasciculation with its major motorneuronal target.

Fenestration nodes and the wide submyelinic space form the basis for the unusually fast impulse conduction of shrimp myelinated axons

Saltatory impulse conduction in invertebrates is rare and has only been found in a few giant nerve fibres, such as the pairs of medial giant fibres with a compact multilayered myelin sheath found in shrimps (Penaeus chinensis and Penaeus japonicus) and the median giant fibre with a loose multilayered myelin sheath found in the earthworm Lumbricus terrestris. Small regions of these nerve fibres are not covered by a myelin sheath and serve as functional nodes for saltatory conduction. Remarkably, shrimp giant nerve fibres have conduction speeds of more than 200 m s-1, making them among the fastest-conducting fibres recorded, even when compared with vertebrate myelinated fibres. A common nodal structure for saltatory conduction has recently been found in the myelinated nerve fibres of the nervous systems of at least six species of Penaeus shrimp, including P. chinensis and P. japonicus. This novel node consists of fenestrated openings that are regularly spaced in the myelin sheath and are designated as fenestration nodes. The myelinated nerve fibres of the Penaeus shrimp also speed impulse conduction by broadening the gap between the axon and the myelin sheath rather than by enlarging the axon diameter as in other invertebrates. In this review, we document and discuss some of the structural and functional characteristics of the myelinated nerve fibres of Penaeus shrimp: (1) the fenestration node, which enables saltatory conduction, (2) a new type of compact multilayered myelin sheath, (3) the unique microtubular sheath that tightly surrounds the axon, (4) the extraordinarily wide space present between the microtubular sheath and the myelin sheath and (5) the main factors contributing to the fastest impulse conduction velocity so far recorded in the Animal Kingdom.

The Orientation of Tail-Flip Escape Swimming in Decapod and Mysid Crustaceans

The orientation of tail-flip escape swimming in a range of adult decapod and mysid crustaceans is reviewed. In mechanical terms, tail-flip swimming constitutes unsteady locomotion in which a single ‘appendage’, the abdomen, produces thrust by a combination of a rowing action and a final ‘squeeze’ force when the abdomen presses against the cephalothorax. In small crustaceans, a symmetrical ‘jack-knife’ tail-flip is more typical. Tail-flip flexion is controlled by two giant-fibre pathways, and also by a non-giant-neuronal network. The direction of thrust in the sagittal plane, and hence the elevation, translation and rotation of the tail-flip are dependent upon the point of stimulation and on the giant-fibre pathway activated. The laterality of the stimulus also affects the orientation of swimming, which is directed away from the point of stimulation. In large decapods such as the lobsters Nephrops norvegicus and Jasus lalandii steering is produced by asym-metrical movements of various abdominal appendages, and by rotation of the abdomen about the cephalothorax. In slipper lobsters the flattened antennae provide steering surfaces. In smaller decapods, such as the brown shrimp Crangon crangon, and in mysids, such as Praunus flexuosus, steering is effected by a rapid rotation of the whole body about its longitudinal axis during the initial stages of tail-flip flexion. The effectiveness of tail-flip swimming is considered in the context of predator-prey interactions under natural conditions and in relation to artificial threats from fishing gear.