Sensory Physiology 275
somatosensory cortex of the postcentral gyrus devoted to the
tongue. Information is also sent to the prefrontal (orbitofron-
tal) cortex, which is important for taste associations and the
perception of flavor.
The specialized epithelial cells of the taste bud are known
as taste cells. The different categories of taste are produced
by different chemicals that come into contact with the micro-
villi of these cells ( fig. 10.8 ). Four different categories of taste
are traditionally recognized: salty, sour, sweet, and bitter.
There is also a more recently discovered fifth category of
taste, termed umami (a Japanese term for “savory,” related to
a meaty flavor), for the amino acid glutamate (and stimulated
by the flavor-enhancer monosodium glutamate). Although sci-
entists long believed that different regions of the tongue were
specialized for different tastes, this is no longer believed to be
true. All areas of the tongue are able to respond to all five cat-
egories of taste. This is true even for a single taste bud, which
can contain taste cells sensitive to each category of taste. How-
ever, a particular taste cell is sensitive to only one category of
taste and activates a sensory neuron that transmits information
regarding that specific taste to the brain.
For example, the sweet taste evoked by sugar is carried to
the brain on sensory neurons devoted only to the sweet taste.
Saccharin in low concentrations stimulates only the sweet
receptors, but at higher concentrations can also stimulate bit-
ter receptors and give saccharine an “aftertaste.” The com-
plex tastes we can perceive depend on the relative activities of
the sensory neurons from each of the five categories of taste,
together with information conveying the sense of smell from
olfactory receptors. Taste is also influenced by the temperature
nucleus of second-order neurons in the medulla oblongata.
From there, the second-order neurons project to the thala-
mus, which serves as a switchboard for directing sensory
information to the cerebral cortex (chapter 8, section 8.2).
Third-order neurons from the thalamus convey taste informa-
tion to the primary gustatory cortex in the insula, and to the
Figure 10.7 A taste bud. Chemicals dissolved in the fluid
at the pore bind to receptor proteins in the microvilli of the sensory
cells. This ultimately leads to the release of neurotransmitter, which
activates the associated sensory neuron.
Sensory
nerve fiber
Supporting
cell
Gustatory
(taste) cell
Taste hair
Taste
bud
Tongue surface Taste pore
Figure 10.8 The five major categories of taste. Each category of taste activates specific taste cells by different means.
Notice that taste cells for salty and sour are depolarized by ions (Na^1 and H^1 , respectively) in the food, whereas taste cells for sweet,
umami, and bitter are depolarized by sugars, the amino acids glutamate and aspartate (not shown), and quinine, respectively, by
means of G-protein-coupled receptors and the actions of second messengers.
x
Na+ H+ Sugars or
amino
acids
Salty Sour Sweet and Umami Bitter
Na+ through ion channel H+ through ion channel
(and other effects)
Binds to membrane receptors
for sweet or umami
Binds to membrane
receptor
Depolarization Depolarization
Depolarization
Opens
Ca2+
channels Ca
2+ released from
endoplasmic
reticulum
Opens
Ca2+
channels
Sensory neuron stimulated Sensory neuron stimulated Sensory neuron stimulated
Neurotransmitter
released
Neurotransmitter
released
Neurotransmitter
released
Neurotransmitter
released
Sensory neuron stimulated
G-proteins
Second
messenger
G-proteins
Second
messenger
Quinine
Close K+
channels
+
+ +
+++
++ ++
+
+
++
Ca2+ Ca2+ Ca2+ Ca2+
+
+
Ca2+