Representations of motivational drives in mesial cortex, medial thalamus, hypothalamus and midbrain.

We propose that neural representations of motivational drives, including sexual desire, hunger, thirst, fear, power-dominance, the motivational aspect of pain, the need for sleep, and nurturance, are represented in four areas in the brain. These are located in the medial hypothalamic/preoptic area, the periaqueductal gray matter (PAG) in the midbrain/pons, the midline and intralaminar thalamic nuclei, and in the anterior part of the mesial cortex, including the medial prefrontal and anterior cingulate areas. We attempt to determine the locations of each of these representations within the hypothalamus/preoptic area, periaqueductal gray and cortex, based on the available literature on activation of brain structures by stimuli that evoke these forms of motivation, on the effects of electrical and chemical stimulation and lesions of candidate structures, and on hodological data. We discuss the hierarchical organization of the representations for a given drive, outputs from these representations to premotor structures in the medulla, caudate-putamen, and cortex, and their contributions to involuntary, learned-sequential (operant) and voluntary behaviors.

Fear and power-dominance drive motivation: neural representations and pathways mediating sensory and mnemonic inputs, and outputs to premotor structures.

Based on the available literature on activation of brain structures by fear- and anger-inducing stimuli, on the effects of electrical and chemical stimulation and lesions of candidate structures, and on connectional data, we propose that both the fear and power-dominance drives are represented in four distinct locations: the medial hypothalamus, lateral/dorsolateral periaqueductal gray, midline thalamic nuclei, and medial prefrontal cortex. The hypothalamic fear representation is located in the dorsomedial and posterior hypothalamic nuclei, the midbrain representation in the caudal part of the lateral/dorsolateral periaqueductal gray, the thalamic representation primarily in parts of the paraventricular and reuniens thalamic nuclei, and the cortical representation in prelimbic cortex. The hypothalamic power-dominance representation is located in the anterior hypothalamic nucleus, dorsomedial aspect of the ventromedial nucleus, and in adjacent parts of the medial preoptic area. The corresponding midbrain representation occurs in rostral part of the lateral/dorsolateral periaqueductal gray, and the thalamic representation in parts of the paraventricular, parataenial, and reuniens thalamic nuclei. We discuss sensory/mnemonic inputs to these representations, and outputs to premotor structures in the medulla, caudate-putamen, and cortex, and their differential contributions to involuntary, learned sequential, and voluntary motor acts. We examine potential contributions of neuronal activities in these representations to the subjective awareness of fear and anger.

Separate, parallel sensory and hedonic pathways in the mammalian somatosensory system.

We propose that separate sensory and hedonic representations exist in each of the primary structures of the somatosensory system, including brainstem, thalamic and cortical components. In the dorsal horn of the spinal cord, the hedonic representation, which consists primarily of nociceptive-specific, wide dynamic range, and thermoreceptive neurons, is located in laminae I and II, while the sensory representation, composed primarily by low-threshold and wide dynamic range neurons, is found in laminae III through V. A similar arrangement is found in the caudal spinal trigeminal nucleus. Based on the available anatomical and electrophysiological data, we then determine the corresponding hedonic and sensory representations in the area of the dorsal column nuclei, ventrobasal and posterior thalamic complex, and cortex. In rodent primary somatosensory cortex, a hedonic representation can be found in laminae Vb and VI. In carnivore and primate primary and secondary somatosensory cortical areas no hedonic representation exists, and the activities of neurons in both areas represent the sensory aspect exclusively. However, there is a hedonic representation in the posterior part of insular cortex, bordering on retroinsular cortex, that receives projections from two thalamic areas in which hedonics are represented. The functions of the segregated components of the system are discussed, especially in relation to the subjective awareness of pain.