in the gray matter of the ventral horn of the spinal cord. In contrast, descending neurons from the reticular formation and motor systems of the cerebral cortex that synapse upon the neurons of the ventral horn serve to modulate their activity and program the sequence of their neuronal discharge patterns, respectively. In a similar manner, the hypothalamus and midbrain PAG constitute the integrating mechanisms whose outputs serve as the "final common pathway" for the expression of aggressive forms of behavior. We propose that the limbic system thus serves the critically important function of modulating the activity of neurons in the hypothalamus and PAG and of programming the sequence of neuronal discharge patterns within these structures. We illustrate this in the examples of research cited below.
Our focus is on two models of aggression that can be elicited by electrical or chemical stimulation of the hypothalamus and midbrain periaqueductal gray matter in the cat—quiet biting "predatory" attack behavior and effective defense behavior. Affective defense behavior may be classified as having aversive properties (such as those associated with fear), whereas predatory attack is associated with more positively reinforcing objectives such as the acquisition of food. The study of these two forms of aggression in the cat may thus provide a more complete picture of the neural substrates of aggression than either model studied alone; moreover, research in the cat has been very systematic and has certain elegant qualities.
Quiet biting attack is predatory in nature and is characterized by stalking of the prey object (usually an anesthetized rat), which is then followed by biting of the back of its neck. The cat may also strike the rat with its forepaw (Flynn et al., 1970; Wasman and Flynn, 1962). This behavior occurs under natural conditions, and includes capturing and killing of a rat or a mouse in the open field (Leyhausen, 1979). In the laboratory, predatory attack behavior can be elicited by electrical stimulation along a region beginning from the rostrolateral and perifornical hypothalamus (Wasman and Flynn, 1962; Siegel and Pott, 1988) and extending caudally through the ventral aspect of the midbrain (Bandler and Flynn, 1972) and ventral aspect of the midbrain PAG (Bandler,
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