Midline lineages in grasshopper produce neuronal siblings with asymmetric expression of Engrailed

The median neuroblast lineage of grasshopper has provided a model for the development of differing neuronal types within the insect central nervous system. According to the prevailing model, neurons of different types are produced in sequence. Contrary to this, we show that each ganglion mother cell from the median neuroblast produces two neurons of asymmetric type: one is Engrailed positive (of interneuronal fate); and one is Engrailed negative (of efferent fate). The mature neuronal population, however, results from differential neuronal death. This yields many interneurons and relatively few efferent neurons. Also contrary to previous reports, we find no evidence for glial production by the median neuroblast. We discuss evidence that neuronal lineages typically produce asymmetric progeny, an outcome that has important developmental and evolutionary implications.