Multiterminal proprioceptors in Squilla mantis (Stomatopoda, Crustacea)

A series of morphologically simple multiterminal proprioceptors is described in the thoracopods of the hoplocaridian crustacean, Squilla mantis, which is a primitive offshoot of the main malacostracan evolutionary line. The description given here is primarily for the largest and most complex receptor that occurs in the second thoracopod, the raptorial limb. These stretch receptors, which lie distally in the coxopodite, consist of single multiterminal sensory neurons with widely branching dendrites innervating broad connective tissue strands. The single primary dendrite divides dichotomously or, occasionally, trichotomously to produce a large number of terminal dendrites. The terminal dendrites, which exhibit a beaded structure, run parallel to the connective tissue strands' longitudinal axis. Dendrites possess an elaborate connective tissue sheath consisting of a single inner glial layer and a multiple outer connective tissue layer. The presence of the beaded structure has been confirmed by transmission electron microscopy (EM), and, because microtubule density does not vary with dendrite diameter, the beads are not fixation artifacts. Beads have high numbers of mitochondria and glycogen deposits, which suggests that they have a high metabolic activity. Dendritic tips are not associated with beads, arise from the terminals of smaller dendrites, and are embedded in the extracellular ground matrix of the connective tissue strand, where they lie parallel to and surround bundles of elastic fibrils. The different degree of development within the receptor series allows comparison between functionally different limbs, and comparison is made with other segmental receptor series. The evolution of multiterminal proprioceptors is discussed.