After synapsing with the mitral cells in the glomeruli of the olfactory bulb, mitral cell axons converge to form the olfactory tract. The olfactory tract then divides into medial and lateral olfactory striae to reach different synaptic targets (eFig. 9.94). Some of the axons in the medial olfactory striae travel through the diagonal band to reach the septal area, whereas others cross the midline in the anterior commissure and inhibit mitral cell activity in the contralateral olfactory bulb to enhance localization of the olfactory stimulant. Axons in the lateral olfactory stria primarily terminate in the piriform cortex/primary olfactory cortex of the uncus and in the amygdala (eFig. 9.94).
The medial forebrain bundle, traveling through the lateral hypothalamus, connects the olfactory cortex with both the hypothalamus and the brainstem to regulate autonomic responses such as arousal through the reticular formation, salivation, and gastric contraction.
The limbic system is composed of several cortical and subcortical structures that participate in an intricate network of connections to regulate complicated behaviors such as memory, emotions, homeostatic functions, and motivational state. In this section we will review the major structures and pathways that form the limbic system.
Grossly, the limbic lobe includes a ring-shaped area of cortical structures that border the brainstem. These cortical areas include the cingulate gyrus, parahippocampal gyrus, and subcallosal area (eFig. 9.12). Laterally, the insular cortex also participates in limbic system function (eFig. 9.10). Nuclear structures of the limbic system include the amygdala, hippocampal formation, anterior and mediodorsal thalamic nuclei, septal nuclei in the forebrain, and nucleus accumbens (eFig. 9.95).
The amygdaloid nucleus is an almond-shaped structure located anterior to the inferior horn of the lateral ventricle and tail of the caudate within the temporal lobe (eFig. 9.96). Structurally, the amygdala consists of three nuclear regions: a large basolateral group and a smaller corticomedial group, which includes the central nucleus. Functionally, the amygdala is primarily associated with the emotion of fear, but it also has an important role in autonomic and neuroendocrine pathways. Connections of the amygdala are predominantly bidirectional and follow three different pathways: the uncinate fasciculus, stria terminalis, and ventral amygdalofugal pathway (eFig. 9.97). Connections to cortical areas pass through the uncinate fasciculus, which progresses anterior to the amygdala. Projections to the septal area and hypothalamus follow the stria terminalis (eFig. 9.98A–D). Fibers forming the ventral amygdalofugal pathway project to thalamic nuclei and several brainstem and forebrain structures (eFig. 9.98A–D).
The nucleus accumbens resides with the ventral forebrain adjacent to where the putamen and head of the caudate become continuous with one another (eFig. 9.99). Afferent axons to the nucleus accumbens come from the amygdala through the amygdalofugal pathway, hippocampal formation by way of the fornix, basal forebrain area from the stria terminalis, and ventral tegmentum through the medial forebrain bundle (eFig. 9.98A–D). Efferent axons leaving the nucleus accumbens project directly to the hypothalamus and globus pallidus and reach nuclei in the brainstem through the medial forebrain bundle. Its connections to the globus pallidus represent an important connection of the limbic system to the motor system. The overall function of the nucleus accumbens is recognized as a gratification center and has been shown to play a role in behaviors related to addiction.