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In several subregions of the mammalian brain, mGluR-mediated PI3K activation is essential for a form of synaptic plasticity termed long-term depression (LTD), which is implicated in neurological diseases such as fragile X and autism. In Drosophila larval motor neurons, ligand activation of DmGluRA, the sole Drosophila mGluR, similarly mediates a PI3K-dependent downregulation of neuronal activity. The mechanism by which mGluR activates PI3K remains incompletely understood in either mammals or Drosophila. Here we identify CaMKII and the nonreceptor tyrosine kinase DFak as critical intermediates in the DmGluRA-dependent activation of PI3K at Drosophila motor nerve terminals. We find that transgene-induced CaMKII inhibition or the DFakCG1 null mutation each block the ability of glutamate application to activate PI3K in larval motor nerve terminals, whereas transgene-induced CaMKII activation increases PI3K activity in motor nerve terminals in a DFak-dependent manner, even in the absence of glutamate application. We also find that CaMKII activation induces other PI3K-dependent effects, such as increased motor axon diameter and increased synapse number at the larval neuromuscular junction. CaMKII, but not PI3K, requires DFak activity for these increases. We conclude that the activation of PI3K by DmGluRA is mediated by CaMKII and DFak.<\/jats:p>","DOI":"10.1534\/genetics.111.128561","type":"journal-article","created":{"date-parts":[[2011,4,23]],"date-time":"2011-04-23T00:34:08Z","timestamp":1303518848000},"page":"601-613","source":"Crossref","is-referenced-by-count":16,"title":["The Metabotropic Glutamate Receptor Activates the Lipid Kinase PI3K in <i>Drosophila<\/i> Motor Neurons Through the Calcium\/Calmodulin-Dependent Protein Kinase II and the Nonreceptor Tyrosine Protein Kinase DFak"],"prefix":"10.1093","volume":"188","author":[{"given":"Curtis","family":"Chun-Jen Lin","sequence":"first","affiliation":[{"name":"Department of Biochemistry and Cell Biology, Rice University, Houston, Texas 77005"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"James B","family":"Summerville","sequence":"additional","affiliation":[{"name":"Department of Biochemistry and Cell Biology, Rice University, Houston, Texas 77005"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Eric","family":"Howlett","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael","family":"Stern","sequence":"additional","affiliation":[{"name":"Department of Biochemistry and Cell Biology, Rice University, Houston, Texas 77005"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"286","published-online":{"date-parts":[[2011,7,1]]},"reference":[{"key":"2022011317224437000_bib1","doi-asserted-by":"crossref","first-page":"123","DOI":"10.1016\/S0898-6568(99)00076-5","article-title":"RAFTK\/Pyk2-mediated cellular signalling","volume":"12","author":"Avraham","year":"2000","journal-title":"Cell Signal"},{"key":"2022011317224437000_bib2","doi-asserted-by":"crossref","first-page":"370","DOI":"10.1016\/j.tins.2004.04.009","article-title":"The mGluR theory of fragile X mental retardation","volume":"27","author":"Bear","year":"2004","journal-title":"Trends Neurosci."},{"key":"2022011317224437000_bib3","doi-asserted-by":"crossref","first-page":"3381","DOI":"10.1242\/dev.129.14.3381","article-title":"Integrins regulate DLG\/FAS2 via a CaM kinase II-dependent pathway to mediate synapse elaboration and stabilization during postembryonic development","volume":"129","author":"Beumer","year":"2002","journal-title":"Development"},{"key":"2022011317224437000_bib4","doi-asserted-by":"crossref","first-page":"9105","DOI":"10.1523\/JNEUROSCI.2724-04.2004","article-title":"The Drosophila metabotropic glutamate receptor DmGluRA regulates activity-dependent synaptic facilitation and fine synaptic morphology","volume":"24","author":"Bogdanik","year":"2004","journal-title":"J. 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