Ventral Median Neurons Research Articles

Independent suboesophageal neuronal innervation of the defense gland and longitudinal muscles in the stick insect (Peruphasma schultei) prothorax

This study investigates the neuroanatomy of the defense gland and a related muscle in the stick insect Peruphasma schultei with axonal tracing and histological sections. The gland is innervated by three neurons through the Nervus anterior of the suboesophageal ganglion (SOG), the ipsilateral neuron (ILN), the contralateral neuron (CLN) and the prothoracic intersegmental neuron (PIN). The ILN has a large soma which is typical for motoneurons that cause fast contraction of large muscles and its dendrites are located in motor-sensory and sensory neuropile areas of the SOG. The CLN might be involved in the coordination of bilateral or unilateral discharge as its neurites are closely associated to the ILN of the contralateral gland. Close to the ejaculatory duct of the gland lies a dorsal longitudinal neck muscle, musculus pronoto-occipitalis (Idlm2), which is likely indirectly involved in gland discharge by controlling neck movements and, therefore, the direction of discharge. This muscle is innervated by three ventral median neurons (VMN). Thus, three neuron types (ILN, CLN, and PIN) innervate the gland muscle directly, and the VMNs could aid secretion indirectly. The cytoanatomy of motorneurons innervating the defense gland and neck muscle are discussed regarding the structure and functions of the neuropile in the SOG. As a basis for the neuroanatomical study on the defense gland we assembled a map of the SOG in Phasmatodea.

Multiple Identified Neurons and Peripheral Nerves Innervating the Prothoracic Defense Glands in Stick Insects Reveal Evolutionary Conserved and Novel Elements of a Chemical Defense System

The defense glands in the dorsal prothorax are an important autapomorphic trait of stick insects (Phasmatodea). Here, we study the functional anatomy and neuronal innervation of the defense glands in Anisomorpha paromalus (Westwood, 1859) (Pseudophasmatinae), a species which sprays its defense secretions when disturbed or attacked. We use a neuroanatomical approach to identify the nerves innervating the gland muscles and the motoneurons with axons in the different nerves. The defense gland is innervated by nerves originating from two segments, the suboesophageal ganglion and the prothoracic ganglion. Axonal tracing confirms the gland innervation via the anterior suboesophageal nerve, and two intersegmental nerves, the posterior suboesophageal nerve and the anterior prothoracic nerve. Axonal tracing of individual nerves reveals eight identified neuron types in the suboesophageal or prothoracic ganglion. The strongest innervating nerve of the gland is the anterior suboesophageal nerve, which also supplies dorsal longitudinal thorax muscles (neck muscles) by separate nerve branches. Tracing of individual nerve branches reveals different sets of motoneurons innervating the defense gland (one ipsilateral and one contralateral suboesophageal neuron) or the neck muscle (ventral median neurons). The ipsilateral and contralateral suboesophageal neurons have no homologs in related taxa like locusts and crickets, and thus evolved within stick insects with the differentiation of the defense glands. The overall innervation pattern suggests that the longitudinal gland muscles derived from dorsal longitudinal neck muscles. In sum, the innervating nerves for dorsal longitudinal muscles are conserved in stick insects, while the neuronal control system was specialized with conserved motoneurons for the persisting neck muscles, and evolutionarily novel suboesophageal and prothoracic motoneurons innervating the defense gland.