Expression of engrailed in an array of identified sensory neurons: comparison with position, axonal arborization, and synaptic connectivity.

engrailed (en) is expressed in the posterior region of embryonic segments and appendages of the cockroach, Periplaneta americana. By 23% of embryogenesis En immunoreactivity is apparent in the dorsal half of the cercus, appendages of segment A11. By 40% of development, En staining is present in the dorsomedial half of the cercus. The nucleus of the medial filiform hair sensory neuron (M), born in this region, expresses en strongly. Staining is never seen in the lateral neuron (L). En is expressed in M as the sensory axons enter the terminal ganglion and begin to form their different arborizations and synaptic connections. This pattern of expression persists through development to the second instar. In mutant animals with supernumerary filiform hair sensilla, En immunoreactivity is only seen in the medial neurons. In second-instar and adult cerci en expression is also seen in medially located neurons. We compared the levels of En staining in the array of 25 second instar neurons with their position, axonal arbor type, and synaptic connections. Staining intensity correlates with distance from the cercal midline, suggesting that en is regulated by other circumferential positional determinants. The expression of en does not correlate with the formation of an M-type arbor. Although 10 to 12 sensory neurons that express en form synapses with giant interneuron 5, the correlation is not precise. These results suggest that, if En does form part of a combinatorial system of positional information in the cercus, its actions are modulated by other gene products.