Glucocorticoids bind to two main types of receptors in the brain, the glucocorticoid receptor (GR) and the mineralocorticoid receptor (MR) (Reul & de Kloet, 1985). Granule cells in the dentate gyrus express both subtypes of adrenal steroid receptors. Because MRs have higher affinity for glucocorticoids than GRs, MRs are more sensitive to circadian changes in glucocorticoids while GRs respond more to stress-induced elevations in glucocorticoids (de Kloet et al., 1998).