Author contributions SC and BLB performed histological analysis. SC, QC, IYMC, HSX, YZ, HM, ZAA, and XD surveyed the intrinsic properties. QC, HMH, XD, YZ, and CJW examined the local connectivity. AP, QC, SR, and NJJ conducted the behavioral studies. IYMC developed the machine learning analysis pipeline. YC and SMB provided expertise for neuron classification. CSC wrote the manuscript with input from all co-authors. All authors reviewed and edited the manuscript. CSC designed, directed, and supervised the project.