The dual human olfactory system. (A) Brain systems involved in smell perception during orthonasal olfaction (sniffing in). (B), Brain systems involved in smell perception during retronasal olfaction (breathing out), with food in the oral cavity. Air flows are indicated by dashed and dotted lines; dotted lines indicate air carrying odor molecules. ACC, accumbens; AM, amygdala; AVI, anterior ventral insular cortex; DI, dorsal insular cortex; LH, lateral hypothalamus; LOFC, lateral orbitofrontal cortex; MOFC, medial orbitofrontal cortex; NST, nucleus of the solitary tract; OB, olfactory bulb; OC, olfactory cortex; OE, olfactory epithelium; PPC, posterior parietal cortex; Pont, pontine taste nucleus; SOM, somatosensory cortex; VII, IX, X, cranial nerves; VC: primary visual cortex; VPM, ventral posteromedial thalamic nucleus. Not shown are numerous additional systems involved in motivation, memory, emotion, reward and language (from Shepherd, 2006).