GPR4 and Anxiety: Additionally, there is significant crossover between efferents and afferents of the GPR4-expressing forebrain nuclei; for example, both the lateral septum and medial habenula project to the dopaminergic neurons of the ventral tegmental area (VTA), whose activation induces anxiety behaviors (Herkenham and Nauta, 1977; Qin and Luo, 2009; Yamaguchi et al., 2013; Anthony et al., 2014; Namboodiri et al., 2016; McLaughlin et al., 2017; Rizzi-Wise and Wang, 2021; Besnard and Leroy, 2022; Gouveia and Ibrahim, 2022; Qi et al., 2022; Wang et al., 2023).