MOUTH-WATERING: THE INFLUENCE OF ENVIRONMENTAL AND COGNITIVE FACTORS ON SALIVATION AND GUSTATORY/FLAVOR PERCEPTION : MAKING THE MOUTH WATER

Oral homeostasis is dependent upon saliva and its content of proteins. Reflex salivary flow occurs at a low 'resting' rate and for short periods of the day more intense taste or chewing stimuli evoke up to ten fold increases in salivation. The secretion of salivary fluid and proteins is controlled by autonomic nerves. All salivary glands are supplied by cholinergic parasympathetic nerves which release acetylcholine that binds to M3 and (to a lesser extent) M1 muscarinic receptors, evoking the secretion of saliva by acinar cells in the endpieces of the salivary gland ductal tree. Most salivary glands also receive a variable innervation from sympathetic nerves which released noradrenaline from which tends to evoke greater release of stored proteins, mostly from acinar cells but also ductal cells. There is some 'cross-talk' between the calcium and cyclic AMP intracellular pathways coupling autonomic stimulation to secretion and salivary protein secretion is augmented during combined stimulation. Other non-adrenergic, non-cholinergic neuropeptides released from autonomic nerves evoke salivary gland secretion and parasympathetically derived vasointestinal peptide, acting through endothelial cell derived nitric oxide, plays a role in the reflex vasodilatation that accompanies secretion. Neuronal type, calcium-activated, soluble nitric oxide within salivary cells appears to play a role in mediating salivary protein secretion in response to autonomimetics. Fluid secretion by salivary glands involves aquaporin 5 and the extent to which the expression of aquaporin 5 on apical acinar cell membranes is upregulated by cholinomimetics remains uncertain. Extended periods of autonomic denervation, liquid diet feeding (reduced reflex stimulation) or duct ligation cause salivary gland atrophy. The latter two are reversible, demonstrating that glands can regenerate provided that the autonomic innervation remains intact. The mechanisms by which nerves integrate with salivary cells during regeneration or during salivary gland development remain to be elucidated.

Following on from ecological theories of perception, such as the one proposed by [Gibson, J. J. (1966). The senses considered as perceptual systems. Boston: Houghton Mifflin] this paper reviews the literature on the multisensory interactions underlying the perception of flavor in order to determine the extent to which it is really appropriate to consider flavor perception as a distinct perceptual system. We propose that the multisensory perception of flavor may be indicative of the fact that the taxonomy currently used to define our senses is simply not appropriate. According to the view outlined here, the act of eating allows the different qualities of foodstuffs to be combined into unified percepts; and flavor can be used as a term to describe the combination of tastes, smells, trigeminal, and tactile sensations as well as the visual and auditory cues, that we perceive when tasting food.