The sensation that develops as a long breath hold continues is what this chapter is about. We term this sensation of an urge to breathe "air hunger". Air hunger is a primal sensation that alerts us to a failure to meet an urgent homeostatic need -to maintain gas exchange. Anxiety, frustration, and fear evoked by air hunger motivate behavioral actions to address the failure. Air hunger can be reliably quantified by most experimental subjects using rating scales, i.e., there is a consistent relationship between stimulus and rating. Stimuli that increase air hunger include hypercapnia, hypoxia, exercise, and acidosis, while tidal expansion of the lungs reduces air hunger. As such, the defining experimental paradigm to evoke air hunger is to elevate the drive to breath while mechanically restricting ventilation. Functional brain imaging studies have shown that air hunger activates the insular cortex (an integration center for perceptions related to homeostasis, including pain, food hunger and thirst), as well as limbic structures involved with anxiety and fear. The unpleasantness and emotional consequences of air hunger make it the most debilitating component of dyspnea, a symptom commonly associated with respiratory, cardiovascular and metabolic diseases. Although much has been learned about air hunger in the past few decades, much remains to be discovered, such as an accepted method to quantify air hunger in non-human animals, fundamental questions about neural mechanisms, and adequate and safe methods to mitigate air hunger in clinical situations.
DIDACTIC SYNOPSIS Air hunger is the conscious appreciation of an uncomfortable urge to breathe. Air hunger can be reliably scaled by most people using various psychophysical rating scales.Although non-human vertebrates presumably sense air hunger, there is currently no accepted experimental model to scale air hunger in animals. Air hunger is a primal sensation, alerting the animal to a threat to homeostasis that requires a behavioral response more complex than increasing respiratory muscle output or cardiac output. Air hunger activates limbic and paralimbic regions in the brain giving rise to anxiety and fear. Air hunger arises when minute ventilation is less than desired minute ventilation. Current understanding is that a copy of motor activity in medullary respiratory centers ('corollary discharge') projects to sensory cortex, and is compared to signals of tidal lung inflation arising mainly in pulmonary stretch receptors. Air hunger is the most uncomfortable and most prominent of the sensations comprising clinical dyspnea.