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Summary Background During development neurons are generated by sequential divisions of neural stem cells, or neuroblasts. In the insect brain progeny of certain stem cells form lineage-specific sets of projections that arborize in distinct brain regions, called clonal units. Though this raises the possibility that the entire neural network in the brain might be organized in a clone-dependent fashion, only a small portion of clones has been identified. Results Using Drosophila melanogaster , we randomly labeled one of about 100 stem cells at the beginning of the larval stage, analyzed the projection patterns of their progeny in the adult, and identified 96 clonal units in the central part of the fly brain, the cerebrum. Neurons of all the clones arborize in distinct regions of the brain, though many clones feature heterogeneous groups of neurons in terms of their projection patterns and neurotransmitters. Arborizations of clones overlap preferentially to form several groups of closely associated clones. Fascicles and commissures were all made by unique sets of clones. Whereas well-investigated brain regions such as the mushroom body and central complex consist of relatively small numbers of clones and are specifically connected with a limited number of neuropils, seemingly disorganized neuropils surrounding them are composed by a much larger number of clones and have extensive specific connections with many other neuropils. Conclusions Our study showed that the insect brain is formed by a composition of cell-lineage-dependent modules. Clonal analysis reveals organized architecture even in those neuropils without obvious structural landmarks.

Summary Background In the fly's visual motion pathways, two cell types—T4 and T5—are the first known relay neurons to signal small-field direction-selective motion responses [1]. These cells then feed into large tangential cells that signal wide-field motion. Recent studies have identified two types of columnar neurons in the second neuropil, or medulla, that relay input to T4 from L1, the ON-channel neuron in the first neuropil, or lamina, thus providing a candidate substrate for the elementary motion detector (EMD) [2]. Interneurons relaying the OFF channel from L1's partner, L2, to T5 are so far not known, however. Results Here we report that multiple types of transmedulla (Tm) neurons provide unexpectedly complex inputs to T5 at their terminals in the third neuropil, or lobula. From the L2 pathway, single-column input comes from Tm1 and Tm2 and multiple-column input from Tm4 cells. Additional input to T5 comes from Tm9, the medulla target of a third lamina interneuron, L3, providing a candidate substrate for L3's combinatorial action with L2 [3]. Most numerous, Tm2 and Tm9's input synapses are spatially segregated on T5's dendritic arbor, providing candidate anatomical substrates for the two arms of a T5 EMD circuit; Tm1 and Tm2 provide a second. Transcript profiling indicates that T5 expresses both nicotinic and muscarinic cholinoceptors, qualifying T5 to receive cholinergic inputs from Tm9 and Tm2, which both express choline acetyltransferase (ChAT). Conclusions We hypothesize that T5 computes small-field motion signals by integrating multiple cholinergic Tm inputs using nicotinic and muscarinic cholinoceptors.