Neural control of vocalization: respiratory and emotional influences

Previous research has shown that a region of the midbrain, the periaqueductal gray matter (PAG), is critical for vocalization. In this review, we describe the results of previous investigations in which we sought to find out how PAG neurons integrate the activity and precise timing of respiratory, laryngeal, and oral muscle activity for natural-sounding vocalization using the technique of excitatory amino acid microinjections in cats. In these studies, all surgical procedures were carried out under deep anaesthesia. In the precollicular decerebrate cat two general types of vocalization, classified as voiced and unvoiced, could be evoked by exciting neurons in the lateral part of the intermediate part of the PAG. The patterns of evoked electromyographic activity were strikingly similar to previously reported patterns of human muscle activity. Coordinated patterns of activity were evoked with just-threshold excitation leading to the conclusion that patterned muscle activity corresponding to the major categories of voiced and voiceless sound production are represented in the PAG. In a parallel series of human and animal experiments, we also determined that the speech and vocalization respiratory patterns are integrated and coordinated with afferent signals related to lung volume. These data have led to the proposal of a new hypothesis for the neural control of vocalization: that the PAG is a crucial brain site for mammalian voice production, not only in the production of emotional or involuntary sounds, but also as a generator of specific respiratory and laryngeal motor patterns essential for human speech and song.