The propodite-dactylopodite (PD) proprioceptor of
Cancer pagurus
consists of a receptor strand in which are embedded the dendrites of up to 80 bipolar nerve cells. The strand is composed of numerous cells and collagen and is surrounded by a thin layer of amorphous connective tissue. Beneath the latter there are often haemocoelic lacunae and individual haemocytes are found surrounded by strand tissue. The sensory cells can be divided into a proximal group with large cell bodies (movement sensitive cells) and a distal row with smaller cell bodies (position sensitive cells). The former can be further divided on the basis of the insertion of their dendrites into the strand. Thus elongation sensitive movement cells (e. s. m. cs) insert into the anterior surface and run in the ventral region of the receptor strand, while relaxation sensitive movement cells (r. s. m. cs) insert into the dorsal surface and run dorsally through the strand. The dendrites run in pairs, the members of each pair being either e. s. m. cs or r. s. m. cs. Each pair is surrounded distally by a scolopidium, consisting of a scolopale cell, two enveloping cells, a tube of extracellular material and, in e. s. m. cs, a canal cell. The scolopale cell contains a scolopale consisting of electron dense material laid down around a matrix of longitudinally oriented microtubules. In e. s. m. cs the enveloping cells also contain aggregations of a similar material. The dendrites make desmosome contacts with the scolopale cell and the corresponding thickenings of the scolopale cell membrane are juxtaposed by scolopale material. Beyond the scolopale the dendrites enter and terminate within a tube of extracellular material. In e. s. m. cs the scolopidium is surrounded mainly by strand cells, but in r. s. m. cs it is surrounded by collagen fibres. Where the collagen fibres meet the sheathing cells there are often contact zones. Other differences between e. s. m. cs and r. s. m. cs occur in the fine structure of the dendrites and these are described. It is suggested that in e. s. m. cs, since the scolopale is apposed by strand cells, stretching of the strand will lead to elongation of the scolopale and hence of the dendritic terminals. In contrast, in r. s. m. cs, it is possible that the scolopidium is not stretched during elongation of the strand, since there is little cellular contact between the two, but that during strand relaxation the energy stored in the collagen fibres forces the proximal end of the scolopale away from the more distal regions and hence stretches the dendritic terminations. Previous theories are discussed and compared with this idea. Also the relative merits of mechanical and physiological explanations for the phasic nature of these units are discussed.