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Among these, multi-member gene families encoding cell-surface proteins are of interest as they can act as neuron-specific recognition tags in combinations and contribute to circuit assembly in complex brains. Recently, a multi-protein interactome has been described between Beat and Side families of IgSF proteins. Here, we use newly generated gene trap transgenic driver lines to probe the spatial expression pattern of\n                    <jats:italic>beat\/side<\/jats:italic>\n                    genes in olfactory receptor neurons (ORNs) and their synaptic target projection neurons (PNs)\n                    <jats:italic>.<\/jats:italic>\n                    Our results revealed that each ORN\/PN class expresses a specific combination of\n                    <jats:italic>beat\/side<\/jats:italic>\n                    genes, hierarchically regulated by lineage-specific genetic programs. To explore whether the class-specific expression of\n                    <jats:italic>beats\/sides<\/jats:italic>\n                    defines ORN-PN matching specificity, we perturbed presynaptic\n                    <jats:italic>beat-IIa<\/jats:italic>\n                    and postsynaptic\n                    <jats:italic>side-IV<\/jats:italic>\n                    in two ORN-PN partners. However, disruption of Beat-IIa-Side-IV interaction did not produce any significant mistargeting in these two examined glomeruli. Our expression mapping revealed that the Beat\/Side interactome between ORNs and PNs appears to be error-tolerant, supporting the robust\n                    <jats:italic>trans-<\/jats:italic>\n                    synaptic recognition. Though without affecting general glomerular targeting, knockdown of\n                    <jats:italic>side<\/jats:italic>\n                    in ORNs leads to the reduction of synaptic development. Interestingly, we found conserved expression patterns of\n                    <jats:italic>beat\/side<\/jats:italic>\n                    orthologs across ORNs in ants and mosquitoes, indicating the shared regulatory strategies specifying the expression of these duplicated paralogs in insect evolution. This also implies the biological significance of\n                    <jats:italic>beats\/sides<\/jats:italic>\n                    in ORN circuit development or function, which is preserved under selective pressure across divergent insect lineages. Overall, this comprehensive analysis of expression patterns lays a foundation for in-depth functional investigations into how Beat\/Side combinatorial expression contributes to the olfactory circuit assembly.\n                  <\/jats:p>","DOI":"10.1371\/journal.pbio.3003955","type":"journal-article","created":{"date-parts":[[2026,8,27]],"date-time":"2026-08-27T20:31:29Z","timestamp":1787862689000},"page":"e3003955","update-policy":"https:\/\/doi.org\/10.1371\/journal.pbio.corrections_policy","source":"Crossref","is-referenced-by-count":0,"title":["Beat- and Side-family cell-surface molecules are expressed combinatorially in the partner neurons of the olfactory circuit in Drosophila"],"prefix":"10.1371","volume":"24","author":[{"given":"Qichen","family":"Duan","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sumie","family":"Okuwa","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Rachel","family":"Estrella","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0004-5629-1297","authenticated-orcid":true,"given":"Chun","family":"Yeung","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Yu-Chieh David","family":"Chen","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Laura Quintana","family":"Rio","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chengcheng","family":"Du","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Khanh M.","family":"Vien","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0001-0626","authenticated-orcid":true,"given":"Pelin Cayirlioglu","family":"Volkan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"340","published-online":{"date-parts":[[2026,8,27]]},"reference":[{"issue":"1","key":"pbio.3003955.ref001","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1534\/genetics.113.154336","article-title":"Genetic control of wiring specificity in the fly olfactory system","volume":"196","author":"W Hong","year":"2014","journal-title":"Genetics"},{"issue":"6","key":"pbio.3003955.ref002","doi-asserted-by":"crossref","first-page":"609","DOI":"10.1002\/wdev.197","article-title":"Mechanisms of olfactory receptor neuron specification in Drosophila","volume":"4","author":"S Barish","year":"2015","journal-title":"Wiley Interdiscip Rev Dev Biol"},{"issue":"1","key":"pbio.3003955.ref003","doi-asserted-by":"crossref","first-page":"50","DOI":"10.1016\/j.semcdb.2005.12.002","article-title":"Wiring specificity in the olfactory system","volume":"17","author":"GSXE Jefferis","year":"2006","journal-title":"Semin Cell Dev Biol"},{"key":"pbio.3003955.ref004","doi-asserted-by":"crossref","first-page":"82","DOI":"10.1007\/978-0-387-78261-4_6","article-title":"Development of the Drosophila olfactory system","volume":"628","author":"V Rodrigues","year":"2008","journal-title":"Adv Exp Med Biol"},{"issue":"1","key":"pbio.3003955.ref005","doi-asserted-by":"crossref","first-page":"85","DOI":"10.1016\/j.conb.2010.11.001","article-title":"Olfactory map formation in the Drosophila brain: genetic specificity and neuronal variability","volume":"21","author":"A Brochtrup","year":"2011","journal-title":"Curr Opin Neurobiol"},{"key":"pbio.3003955.ref006","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.72599","article-title":"Chemoreceptor co-expression in Drosophila melanogaster olfactory neurons","volume":"11","author":"D Task","year":"2022","journal-title":"Elife"},{"issue":"6860","key":"pbio.3003955.ref007","doi-asserted-by":"crossref","first-page":"204","DOI":"10.1038\/35102574","article-title":"Target neuron prespecification in the olfactory map of Drosophila","volume":"414","author":"GS Jefferis","year":"2001","journal-title":"Nature"},{"issue":"8","key":"pbio.3003955.ref008","article-title":"A complete developmental sequence of a Drosophila neuronal lineage as revealed by twin-spot MARCM","volume":"8","author":"H-H Yu","year":"2010","journal-title":"PLoS Biol"},{"issue":"1","key":"pbio.3003955.ref009","doi-asserted-by":"crossref","first-page":"5","DOI":"10.1186\/s13064-018-0102-0","article-title":"Linking neuronal lineage and wiring specificity","volume":"13","author":"H Li","year":"2018","journal-title":"Neural Dev"},{"issue":"8","key":"pbio.3003955.ref010","doi-asserted-by":"crossref","DOI":"10.1101\/cshperspect.a001776","article-title":"Topographic mapping--the olfactory system","volume":"2","author":"T Imai","year":"2010","journal-title":"Cold Spring Harb Perspect Biol"},{"issue":"2","key":"pbio.3003955.ref011","first-page":"157","article-title":"From lineage to wiring specificity. 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