The Arf6 activator Efa6/PSD3 confers regional specificity and modulates ethanol consumption in Drosophila and humans


Ubiquitously expressed genes have been implicated in a variety of specific behaviors, including responses to ethanol. However, the mechanisms that confer this behavioral specificity have remained elusive. Previously, we showed that the ubiquitously expressed small GTPase Arf6 is required for normal ethanol-induced sedation in adult Drosophila. Here, we show that this behavioral response also requires Efa6, one of (at least) three Drosophila Arf6 guanine exchange factors. Ethanol-naive Arf6 and Efa6 mutants were sensitive to ethanol-induced sedation and lacked rapid tolerance upon re-exposure to ethanol, when compared with wild-type flies. In contrast to wild-type flies, both Arf6 and Efa6 mutants preferred alcohol-containing food without prior ethanol experience. An analysis of the human ortholog of Arf6 and orthologs of Efa6 (PSD1-4) revealed that the minor G allele of single nucleotide polymorphism (SNP) rs13265422 in PSD3, as well as a haplotype containing rs13265422, was associated with an increased frequency of drinking and binge drinking episodes in adolescents. The same haplotype was also associated with increased alcohol dependence in an independent European cohort. Unlike the ubiquitously expressed human Arf6 GTPase, PSD3 localization is restricted to the brain, particularly the prefrontal cortex (PFC). Functional magnetic resonance imaging revealed that the same PSD3 haplotype was also associated with a differential functional magnetic resonance imaging signal in the PFC during a Go/No-Go task, which engages PFC-mediated executive control. Our translational analysis, therefore, suggests that PSD3 confers regional specificity to ubiquitous Arf6 in the PFC to modulate human alcohol-drinking behaviors.

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We thank the Bloomington stock center, and Yang Hong (University of Pittsburgh) for fly strains, and Michael Buszczak, Helmut Krämer and Rothenfluh lab members for helpful discussions. This work was supported by the NIH (T32 fellowships DA007290 to DAG and JHP, F32 AA021340 to SAO, K08 DK091316 to ARR, R01 AA019526 and R21 AA022404 to AR), the European Union-funded FP6 Integrated Project IMAGEN (Reinforcement-related behavior in normal brain function and psychopathology; LSHM-CT-2007–037286), the FP7 projects IMAGEMEND (602450) and MATRICS (603016), the Innovative Medicine Initiative Project EU-AIMS (115300-2), the European Research Council Award ‘STRATIFY’ as well as the Medical Research Council Programme Grant ‘Developmental pathways into adolescent substance abuse’ (93558). Further support was provided by the Swedish Funding Agency FORMAS, the MRC-ICMR Newton project ‘Consortium on Vulnerability to Externalizing Disorders and Addictions’ (c-VEDA) (MR/N000390/1), the National Institute for Health Research (NIHR) Biomedical Research Centre at South London and Maudsley NHS Foundation Trust and King’s College London, the German Bundesministerium für Bildung und Forschung (BMBF grants 01GS08152; 01EV0711; eMED SysAlc 01ZX1311A; Forschungsnetz AERIAL), the NIH (R01 MH085772-01A1), as well as the NIH-BD2K (Big Data to Knowledge) grant U54 EB020403–ENIGMA Center for Worldwide Medicine, Imaging and Genomics. AR was also supported by an Effie Marie Cain Scholarship in Biomedical Research from UT Southwestern Medical Center Dallas.

Author contributions

DAG, JHP, SAO, SFA and AR conceived, performed and analyzed the Drosophila experiments. SD, TB, CB, ALWB, PJC, HF, BI, ML, JM, TP, MS, GS and the IMAGEN consortium acquired the human data. TJ, BX and GS analyzed the human data. DAG, TJ, JHP, JLH, ARR, GS and AR wrote the paper.

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Correspondence to G Schumann or A Rothenfluh.

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Competing interests

TB served in an advisory or consultancy role for Hexal Pharma, Lilly, Medice, Novartis, Otsuka, Oxford Outcomes, PCM Scientific, Shire and Viforpharma. TB received conference attendance support and conference support or speaker’s fees from Lilly, Medice, Novartis and Shire. TB is/has been involved in clinical trials conducted by Shire and Viforpharma. JG has received research funding from AstraZeneca, Eli Lilly & Co., Janssen-Cilag and Bristol-Myers Squibb, and speaker’s fees from AstraZeneca, Janssen-Cilag and Bristol-Myers Squibb. The remaining authors declare no conflict of interest.

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Gonzalez, D., Jia, T., Pinzón, J. et al. The Arf6 activator Efa6/PSD3 confers regional specificity and modulates ethanol consumption in Drosophila and humans. Mol Psychiatry 23, 621–628 (2018).

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