Aftertaste
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Aftertaste is the
taste The gustatory system or sense of taste is the sensory system that is partially responsible for the perception of taste (flavor). Taste is the perception produced or stimulated when a substance in the mouth reacts chemically with taste receptor ...
intensity of a
food Food is any substance consumed by an organism for nutritional support. Food is usually of plant, animal, or fungal origin, and contains essential nutrients, such as carbohydrates, fats, proteins, vitamins, or minerals. The substance is inge ...
or
beverage A drink or beverage is a liquid intended for human consumption. In addition to their basic function of satisfying thirst, drinks play important roles in human culture. Common types of drinks include plain drinking water, milk, juice, smoothies a ...
that is perceived immediately after that food or beverage is removed from the mouth. The aftertastes of different foods and beverages can vary by intensity and over time, but the unifying feature of aftertaste is that it is perceived ''after'' a food or beverage is either swallowed or spat out. The
neurobiological Neuroscience is the scientific study of the nervous system (the brain, spinal cord, and peripheral nervous system), its functions and disorders. It is a multidisciplinary science that combines physiology, anatomy, molecular biology, developme ...
mechanisms of taste (and aftertaste)
signal transduction Signal transduction is the process by which a chemical or physical signal is transmitted through a cell as a series of molecular events, most commonly protein phosphorylation catalyzed by protein kinases, which ultimately results in a cellula ...
from the taste receptors in the mouth to the brain have not yet been fully understood. However, the primary taste processing area located in the insula has been observed to be involved in aftertaste perception.


Temporal taste perception

Characteristics of a food's aftertaste are quality, intensity, and duration. Quality describes the actual taste of a food and intensity conveys the magnitude of that taste. Duration describes how long a food's aftertaste sensation lasts. Foods that have lingering aftertastes typically have long sensation durations. Because taste perception is unique to every person, descriptors for taste quality and intensity have been standardized, particularly for use in scientific studies. For taste quality, foods can be described by the commonly used terms "sweet", "sour", "salty", "bitter", "umami", or "no taste". Description of aftertaste perception relies heavily upon the use of these words to convey the taste that is being sensed after a food has been removed from the mouth. The description of taste intensity is also subject to variability among individuals. Variations of the Borg Category Ratio Scale or other similar metrics are often used to assess the intensities of foods. The scales typically have categories that range from either zero or one through ten (or sometimes beyond ten) that describe the taste intensity of a food. A score of zero or one would correspond to unnoticeable or weak taste intensities, while a higher score would correspond to moderate or strong taste intensities. It is the prolonged moderate or strong taste intensities that persist even after a food is no longer present in the mouth that describe aftertaste sensation. Foods that have distinct aftertastes are distinguished by their temporal profiles, or how long their tastes are perceived during and after consumption. A sample testing procedure to measure a food's temporal profile would entail first recording the time of onset for initial taste perception when the food is consumed, and then recording the time at which there is no longer any perceived taste. The difference between these two values yields the total time of taste perception. Match this with intensity assessments over the same time interval and a representation of the food's taste intensity over time can be obtained. With respect to aftertaste, this type of testing would have to measure the onset of taste perception from the point after which the food was removed from the mouth.


Variability of human taste perception

The categorization of people into "tasters" or "nontasters" - based on their sensitivity to the bitterness of
propylthiouracil Propylthiouracil (PTU) is a medication used to treat hyperthyroidism. This includes hyperthyroidism due to Graves' disease and toxic multinodular goiter. In a thyrotoxic crisis it is generally more effective than methimazole. Otherwise it is t ...
and the expression of
fungiform papillae Lingual papillae (singular papilla) are small structures on the upper surface of the tongue that give it its characteristic rough texture. The four types of papillae on the human tongue have different structures and are accordingly classified as c ...
on their tongues - has suggested a genetic basis for the variations observed in taste perception from person to person. This might imply that the activities of specific genes that affect an individual's perception of different foods' sensations of aftertaste could also affect an individual's perception of different foods. For example, the intensity of the aftertaste sensations "nontasters" experienced after caffeine consumption was found to diminish faster than the sensations "tasters" experienced. This may imply that because of their taste-bud profiles, "tasters" may be more sensitive to the tastes of different foods, and thus experience a more persistent sensation of those foods' tastes.


Taste receptor dynamics

Because a lingering taste sensation is intrinsic to aftertaste, the molecular mechanisms that underlie aftertaste are presumed to be linked to either the continued or delayed activation of receptors and signaling pathways in the mouth that are involved in taste processing. The current understanding of how a food's taste is communicated to the brain is as follows: # Chemicals in food interact with receptors on the taste receptor cells located on the tongue and the roof of the mouth. These interactions can be affected by temporal and spatial factors like the time of receptor activation or the particular taste receptors that are activated (sweet, salty, bitter, etc.). # The
chorda tympani The chorda tympani is a branch of the facial nerve that originates from the taste buds in the front of the tongue, runs through the middle ear, and carries taste messages to the brain. It joins the facial nerve (cranial nerve VII) inside the fa ...
(cranial nerve VII), the
glossopharyngeal nerve The glossopharyngeal nerve (), also known as the ninth cranial nerve, cranial nerve IX, or simply CN IX, is a cranial nerve that exits the brainstem from the sides of the upper Medulla oblongata, medulla, just anterior (closer to the nose) to t ...
(cranial nerve IX), and the
vagus nerve The vagus nerve, also known as the tenth cranial nerve, cranial nerve X, or simply CN X, is a cranial nerve that interfaces with the parasympathetic control of the heart, lungs, and digestive tract. It comprises two nerves—the left and right ...
(cranial nerve X) carry information from the taste receptors to the brain for cortical processing. In the context of aftertaste, the combination of both receptor-dependent and receptor-independent processes have been proposed to explain the signal transduction mechanisms for foods with distinct aftertastes, particularly those that are bitter. The receptor-dependent process is the same as what was described above. However, the receptor-independent process involves the diffusion of bitter,
amphiphilic An amphiphile (from the Greek αμφις amphis, both, and φιλíα philia, love, friendship), or amphipath, is a chemical compound possessing both hydrophilic (''water-loving'', polar) and lipophilic (''fat-loving'') properties. Such a compo ...
chemicals like
quinine Quinine is a medication used to treat malaria and babesiosis. This includes the treatment of malaria due to ''Plasmodium falciparum'' that is resistant to chloroquine when artesunate is not available. While sometimes used for nocturnal leg cr ...
across the taste receptor cell membranes. Once inside the taste receptor cell, these compounds have been observed to activate intracellular
G-proteins G proteins, also known as guanine nucleotide-binding proteins, are a family of proteins that act as molecular switches inside cells, and are involved in transmitting signals from a variety of stimuli outside a cell to its interior. Their ac ...
and other proteins that are involved in signaling pathways routed to the brain. The bitter compounds thus activate both the taste receptors on the cell surface, as well as the signaling pathway proteins in the
intracellular space Intracellular space is the interior space of the plasma membrane. It contains about two-thirds of TBW. Cellular rupture may occur if the intracellular space becomes dehydrated, or if the opposite happens, where it becomes too bloated. Thus it ...
. Intracellular signaling may be slower than taste cell receptor activation since more time is necessary for the bitter compounds to diffuse across the cell membrane and interact with intracellular proteins. This delayed activation of intracellular signaling proteins in response to the bitter compounds, in addition to the extracellular receptor signaling is proposed to be related to the lingering aftertaste associated with bitter foods. The combination of both mechanisms leads to an overall longer response of the taste receptor cells to the bitter foods, and aftertaste perception subsequently occurs.


Processing in the cerebral cortex

The primary taste perception areas in the cerebral cortex are located in the insula and regions of the
somatosensory cortex In physiology, the somatosensory system is the network of neural structures in the brain and body that produce the perception of touch (haptic perception), as well as temperature (thermoception), body position (proprioception), and pain. It is ...
; the
nucleus of the solitary tract In the human brainstem, the solitary nucleus, also called nucleus of the solitary tract, nucleus solitarius, and nucleus tractus solitarii, (SN or NTS) is a series of purely sensory nuclei (clusters of nerve cell bodies) forming a vertical column ...
located in the brainstem also plays a major role in taste perception. These regions were identified when human subjects were exposed to a taste stimulus and their cerebral blood flow measured with
magnetic resonance imaging Magnetic resonance imaging (MRI) is a medical imaging technique used in radiology to form pictures of the anatomy and the physiological processes of the body. MRI scanners use strong magnetic fields, magnetic field gradients, and radio wave ...
. Although these regions have been identified as the primary zones for taste processing in the brain, other cortical areas are also activated during eating, as other sensory inputs are being signaled to the cortex. For aftertaste, much is unclear about the cortical processing related to its perception. The first neuroimaging study to evaluate the temporal taste profile of
aspartame Aspartame is an artificial non-saccharide sweetener 200 times sweeter than sucrose and is commonly used as a sugar substitute in foods and beverages. It is a methyl ester of the aspartic acid/phenylalanine dipeptide with the trade names ...
, an artificial sweetener, in humans was published in 2009. In it, the insula was observed to be activated for a longer period of time than other sensory processing areas in the brain when the aftertaste profile of aspartame was measured. Subjects were administered a solution of aspartame for a specific amount of time before being instructed to swallow the solution. Functional magnetic resonance images of the blood flow in the subjects' brains were recorded before and after they swallowed the aspartame solution. Before swallowing, the
amygdala The amygdala (; plural: amygdalae or amygdalas; also '; Latin from Greek, , ', 'almond', 'tonsil') is one of two almond-shaped clusters of nuclei located deep and medially within the temporal lobes of the brain's cerebrum in complex verteb ...
, somatosensory cortex,
thalamus The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...
, and
basal ganglia The basal ganglia (BG), or basal nuclei, are a group of subcortical nuclei, of varied origin, in the brains of vertebrates. In humans, and some primates, there are some differences, mainly in the division of the globus pallidus into an extern ...
were all activated. After swallowing, only the insula remained activated and the response of the other brain regions was not evident. This suggests that the insula may be a primary region for aftertaste sensation because it was activated even after the aspartame solution was no longer present in the mouth. This finding aligns with the insula's identification as a central taste processing area and simply expands its function. An explanation for less activation of the amygdala was that because it is a reward center in the brain, less reward would be experienced by the subjects during prolonged exposure to the aspartame solution.


Distinguishing aftertaste and flavor

Flavor Flavor or flavour is either the sensory perception of taste or smell, or a flavoring in food that produces such perception. Flavor or flavour may also refer to: Science *Flavors (programming language), an early object-oriented extension to Lis ...
is an emergent property that is the combination of multiple sensory systems including
olfaction The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, it ...
, taste, and somatosensation.Stevenson, R.J. (2009). The psychology of flavour. Oxford: Oxford University Press. How the flavor of a food is perceived, whether it is unpleasant or satisfying, is stored as a memory so that the next time the same (or a similar) food is encountered, the previous experience can be recalled and a decision made to consume that food. This process of multisensory inputs to the brain during eating, followed by learning from eating experiences is the central idea of flavor processing. Richard Stevenson mentions in ''The Psychology of Flavour'' that people often do not realize that a food's flavor can be described by the food's smell, taste, or texture. Instead, he claims, people perceive flavor as a "unitary percept", in which a descriptor for either taste or smell is used to describe a food's flavor. Consider the terms that are used to describe the flavors of foods. For instance, a food may ''taste'' sweet, but often its flavor is described as such while not considering its smell or other sensory characteristics. For example, honey ''tastes'' sweet so its smell is associated with that descriptor, and ''sweet'' is also used to describe its flavor. In fact, sweetness is one of the four basic taste qualities and only comprises part of a food's flavor. Unlike flavor, aftertaste is a solely gustatory event that is not considered to involve any of the other major senses. The distinction of being based on one (aftertaste) versus multiple (flavor) sensory inputs is what separates the two phenomena.


Foods with distinct aftertastes


Artificial sweeteners

Low-calorie artificial sweeteners like
saccharin Saccharin (''aka'' saccharine, Sodium sacchari) is an artificial sweetener with effectively no nutritional value. It is about 550 times as sweet as sucrose but has a bitter or metallic aftertaste, especially at high concentrations. Saccharin is u ...
and acesulfame-K are known for their bitter aftertastes. Recently, GIV3727 (4-(2,2,3-trimethylcyclopentyl) butanoic acid), a chemical that blocks saccharin and acesulfame-K activation of multiple bitter
taste receptor A taste receptor or tastant is a type of cellular receptor which facilitates the sensation of taste. When food or other substances enter the mouth, molecules interact with saliva and are bound to taste receptors in the oral cavity and other locat ...
s has been developed. In the study, the addition of the bitter taste
receptor antagonist A receptor antagonist is a type of receptor ligand or drug that blocks or dampens a biological response by binding to and blocking a receptor rather than activating it like an agonist. Antagonist drugs interfere in the natural operation of rece ...
GIV3727 to the saccharin and acesulfame-K solutions resulted in significantly lower taste intensity ratings when compared to the solutions that were not treated with GIV3727. This suggests that GIV3727 inhibits the normal functions of the bitter taste receptors because saccharin and acesulfame-K's bitter aftertastes were not observed. The ability to inhibit activation of the bitter taste receptors can have far-reaching effects if the bitter aftertastes of not only these two artificial sweeteners but also other foods, beverages, and even pharmaceuticals can be minimized.


Wine

In
wine tasting Wine tasting is the sensory examination and evaluation of wine. While the practice of wine tasting is as ancient as its production, a more formalized methodology has slowly become established from the 14th century onward. Modern, professional w ...
the aftertaste or ''finish'' of a wine, is an important part of the evaluation.Stone, H, & L., J. (2004). Sensory evaluation practices. Amsterdam: Elsevier Academic Press. After tasting a wine, a taster will determine the wine's aftertaste, which is a major determinant of the wine's quality. The aftertaste of a wine can be described as bitter, persistent, short, sweet, smooth, or even non-existent. Included in assessing the aftertaste of a wine is consideration of the aromas still present after swallowing. High quality wines typically have long finishes accompanied by pleasant aromas. By assessing the combination of olfactory and aftertaste sensations, wine tasting actually determines not only the aftertaste profile of a wine, but its flavor profile as well.


References

{{Taste Sensory systems Gustation Gustatory system