Gustatory cortex
Encyclopedia
The primary gustatory cortex is a brain structure responsible for the perception of taste
. It consists of two substructures: the anterior insula on the insular lobe and the frontal operculum on the inferior frontal gyrus of the frontal lobe. Because of its composition the primary gustatory cortex is sometimes referred to in literature as the AI/FO(Anterior Insula/Frontal Operculum). By using extracellular unit recording techniques, scientists have elucidated that neurons in the AI/FO respond to sweetness, saltiness, bitterness, and sourness, and they code the intensity of the taste stimulus.
, the taste system is defined by its specialized peripheral receptors and central pathways that relay and process taste information. Peripheral taste receptors are found on the upper surface of the tongue, soft palate, pharynx
, and the upper part of the esophagus
. Taste cells synapse with primary sensory axons that run in the chorda tympani and greater superficial petrosal branches of the facial nerve
(cranial nerve VII), the lingual branch of the glossopharyngeal nerve
(cranial nerve IX), and the superior laryngeal branch of the vagus nerve
(Cranial nerve X) to innervate the taste buds in the tongue, palate, epiglottis
, and esophagus respectively. The central axons of these primary sensory neurons in the respective cranial nerve ganglia project to rostral and lateral regions of the nucleus of the solitary tract in the medulla
, which is also known as the gustatory nucleus of the solitary tract complex. Axons from the rostral (gustatory) part of the solitary nucleus project to the ventral posterior complex of the thalamus
, where they terminate in the medial half of the ventral posterior medial nucleus. This nucleus projects in turn to several regions of the neocortex
which includes the gustatory cortex (the frontal operculum and the insula), which becomes activated when the subject is consuming and experiencing taste.
, chorda tympani
, and a lingual branch of the glossopharyngeal nerve
elicit evoked field potential in the frontal operculum. Electrical stimulation of the insula in the human elicit gustatory sensations. Gustatory information is conveyed to the orbitofrontal cortex, the secondary gustatory cortex from the AI/FO. Studies have shown that 7.9% of neurons in the orbitofrontal cortex respond to taste stimuli, and a part of these neurons are finely tuned to particular taste stimuli. It has also been shown in monkeys that the responses of orbitofrontal neurons to taste decreased when the monkey eats to satiety. Furthermore neurons in the orbitofrontal cortex respond to the visual ,and/or olfactory stimuli in addition to the gustatory stimulus. These results suggest that gustatory neurons in the orbitofrontal cortex may play an important role in food identification and selection. A patient study reported that damage in the rostral part of the insula caused gustatory disturbance, as well as taste recognition and intensity deficits in patients with insular cortex lesions. It has also been reported that a patient who had an epileptic focus in the frontal operculum and epileptic activity in the focus produced a disagreeable taste. Activation in the insula also takes place when exposed to gustatory imagery. Studies compared the activated regions in subjects shown food pictures to those shown location pictures and found that food pictures activated the right insula/operculum and the left orbitofrontal cortex.
) concentration lingual exposure results in an increase in firing rate in the rat GC neurons, whereas an increase in sucrose concentration resulted in a decrease in firing rate. GC neurons exhibit rapid and selective response to tastants. Sodium Chloride and Sucrose elicited the largest response in the rat gustatory cortex in rats, whereas citric acid causes only a moderate increase in activity in a single neuron. Chemosensory GC neurons are broadly tuned, meaning that a larger percentage of them respond to larger amount of tastants (4 and 5) as compared to the lower percentage responding to a fewer amount of tastants (1 and 2). In addition the number of neurons responding to a certain tastant stimuli varies. In the rat gustatory complex study it was shown that more neurons responded to MSG, NaCl, sucrose
, and citric acid (all activating approximately the same percentage of neurons) as compared to the compounds quinine (QHCl) and water.
), the vast majority of them responded to concentration changes in a complex manner. In such instances with several concentration tastants tested, the middle concentration might evoke the highest firing rate (like 0.1 M sucrose), or the highest and lowest concentrations might elicit the highest rates (NaCl ), or the neuron might respond to only one concentration.
receptors, modulation of gene expression, and posttranslational modifications detected in the insular cortex in the first hours after the consumption of an unfamiliar tastant.
Taste
Taste is one of the traditional five senses. It refers to the ability to detect the flavor of substances such as food, certain minerals, and poisons, etc....
. It consists of two substructures: the anterior insula on the insular lobe and the frontal operculum on the inferior frontal gyrus of the frontal lobe. Because of its composition the primary gustatory cortex is sometimes referred to in literature as the AI/FO(Anterior Insula/Frontal Operculum). By using extracellular unit recording techniques, scientists have elucidated that neurons in the AI/FO respond to sweetness, saltiness, bitterness, and sourness, and they code the intensity of the taste stimulus.
The Taste Pathway and The Role of Gustatory Cortex (GC)
Like the olfactory systemOlfactory system
The olfactory system is the sensory system used for olfaction, or the sense of smell. Most mammals and reptiles have two distinct parts to their olfactory system: a main olfactory system and an accessory olfactory system. The main olfactory system detects volatile, airborne substances, while the...
, the taste system is defined by its specialized peripheral receptors and central pathways that relay and process taste information. Peripheral taste receptors are found on the upper surface of the tongue, soft palate, pharynx
Pharynx
The human pharynx is the part of the throat situated immediately posterior to the mouth and nasal cavity, and anterior to the esophagus and larynx. The human pharynx is conventionally divided into three sections: the nasopharynx , the oropharynx , and the laryngopharynx...
, and the upper part of the esophagus
Esophagus
The esophagus is an organ in vertebrates which consists of a muscular tube through which food passes from the pharynx to the stomach. During swallowing, food passes from the mouth through the pharynx into the esophagus and travels via peristalsis to the stomach...
. Taste cells synapse with primary sensory axons that run in the chorda tympani and greater superficial petrosal branches of the facial nerve
Facial nerve
The facial nerve is the seventh of twelve paired cranial nerves. It emerges from the brainstem between the pons and the medulla, and controls the muscles of facial expression, and functions in the conveyance of taste sensations from the anterior two-thirds of the tongue and oral cavity...
(cranial nerve VII), the lingual branch of the glossopharyngeal nerve
Glossopharyngeal nerve
The glossopharyngeal nerve is the ninth of twelve pairs of cranial nerves . It exits the brainstem out from the sides of the upper medulla, just rostral to the vagus nerve...
(cranial nerve IX), and the superior laryngeal branch of the vagus nerve
Vagus nerve
The vagus nerve , also called pneumogastric nerve or cranial nerve X, is the tenth of twelve paired cranial nerves...
(Cranial nerve X) to innervate the taste buds in the tongue, palate, epiglottis
Epiglottis
The epiglottis is a flap that is made of elastic cartilage tissue covered with a mucous membrane, attached to the entrance of the larynx. It projects obliquely upwards behind the tongue and the hyoid bone, pointing dorsally. The term, like tonsils, is often incorrectly used to refer to the uvula...
, and esophagus respectively. The central axons of these primary sensory neurons in the respective cranial nerve ganglia project to rostral and lateral regions of the nucleus of the solitary tract in the medulla
Medulla oblongata
The medulla oblongata is the lower half of the brainstem. In discussions of neurology and similar contexts where no ambiguity will result, it is often referred to as simply the medulla...
, which is also known as the gustatory nucleus of the solitary tract complex. Axons from the rostral (gustatory) part of the solitary nucleus project to the ventral posterior complex of the thalamus
Thalamus
The thalamus is a midline paired symmetrical structure within the brains of vertebrates, including humans. It is situated between the cerebral cortex and midbrain, both in terms of location and neurological connections...
, where they terminate in the medial half of the ventral posterior medial nucleus. This nucleus projects in turn to several regions of the neocortex
Neocortex
The neocortex , also called the neopallium and isocortex , is a part of the brain of mammals. It is the outer layer of the cerebral hemispheres, and made up of six layers, labelled I to VI...
which includes the gustatory cortex (the frontal operculum and the insula), which becomes activated when the subject is consuming and experiencing taste.
Functionality and Stimulation of Gustatory Cortex (GC)
There have been many studies done to observe the functionality of the primary gustatory cortex and associated structures with various chemical and electrical stimulations as well as observations of patients with lesions and GC epileptic focus. It has been reported that electrical stimulation of the lingual nerveLingual nerve
The lingual nerve is a branch of the mandibular nerve , itself a branch of the trigeminal nerve, which supplies sensory innervation to the tongue...
, chorda tympani
Chorda tympani
The chorda tympani is a nerve that branches from the facial nerve inside the facial canal, just before the facial nerve exits the skull via the Stylomastoid foramen...
, and a lingual branch of the glossopharyngeal nerve
Glossopharyngeal nerve
The glossopharyngeal nerve is the ninth of twelve pairs of cranial nerves . It exits the brainstem out from the sides of the upper medulla, just rostral to the vagus nerve...
elicit evoked field potential in the frontal operculum. Electrical stimulation of the insula in the human elicit gustatory sensations. Gustatory information is conveyed to the orbitofrontal cortex, the secondary gustatory cortex from the AI/FO. Studies have shown that 7.9% of neurons in the orbitofrontal cortex respond to taste stimuli, and a part of these neurons are finely tuned to particular taste stimuli. It has also been shown in monkeys that the responses of orbitofrontal neurons to taste decreased when the monkey eats to satiety. Furthermore neurons in the orbitofrontal cortex respond to the visual ,and/or olfactory stimuli in addition to the gustatory stimulus. These results suggest that gustatory neurons in the orbitofrontal cortex may play an important role in food identification and selection. A patient study reported that damage in the rostral part of the insula caused gustatory disturbance, as well as taste recognition and intensity deficits in patients with insular cortex lesions. It has also been reported that a patient who had an epileptic focus in the frontal operculum and epileptic activity in the focus produced a disagreeable taste. Activation in the insula also takes place when exposed to gustatory imagery. Studies compared the activated regions in subjects shown food pictures to those shown location pictures and found that food pictures activated the right insula/operculum and the left orbitofrontal cortex.
Chemosensory Neurons of the Gustatory Cortex
Chemosensory neurons are defined are those that discriminate between tastant as well as between the presence or absence of a tastant. In these neurons, the responses to reinforced (stimulated by tastant) licks in rats were greater than to those for the unreinforced (not stimulated by tastant) licks. They found that 34.2% of the GC neurons exhibited chemosensory responses. The remaining neurons discriminate between reinforced and unreinforced licks, or process task related information.Tastant Concentration-dependent neuronal activity
GC chemosensory neurons exhibit concentration dependent responses. In a study done on GC responses in rats during licking, an increase in MSG (Monosodium GlutamateMonosodium glutamate
Monosodium glutamate, also known as sodium glutamate or MSG, is the sodium salt of glutamic acid, one of the most abundant naturally occurring non-essential amino acids....
) concentration lingual exposure results in an increase in firing rate in the rat GC neurons, whereas an increase in sucrose concentration resulted in a decrease in firing rate. GC neurons exhibit rapid and selective response to tastants. Sodium Chloride and Sucrose elicited the largest response in the rat gustatory cortex in rats, whereas citric acid causes only a moderate increase in activity in a single neuron. Chemosensory GC neurons are broadly tuned, meaning that a larger percentage of them respond to larger amount of tastants (4 and 5) as compared to the lower percentage responding to a fewer amount of tastants (1 and 2). In addition the number of neurons responding to a certain tastant stimuli varies. In the rat gustatory complex study it was shown that more neurons responded to MSG, NaCl, sucrose
Sucrose
Sucrose is the organic compound commonly known as table sugar and sometimes called saccharose. A white, odorless, crystalline powder with a sweet taste, it is best known for its role in human nutrition. The molecule is a disaccharide composed of glucose and fructose with the molecular formula...
, and citric acid (all activating approximately the same percentage of neurons) as compared to the compounds quinine (QHCl) and water.
Responsiveness of GC neurons to changes in concentration
Studies using the Gustatory cortex of the rat model have shown that GC neurons exhibit complex responses to changes in concentration of tastant. For one tastant, the same neuron might increase its firing rate where as for another tastant, it may only be responsive to an intermediate concentration. Studies have shown that few chemosensory GC neurons. In these studies it was evident that few chemosensory GC neurons monotonically increased or decreased their firing rates in response to changes in concentration of tastants (such as MSG, NaCl, and SucroseSucrose
Sucrose is the organic compound commonly known as table sugar and sometimes called saccharose. A white, odorless, crystalline powder with a sweet taste, it is best known for its role in human nutrition. The molecule is a disaccharide composed of glucose and fructose with the molecular formula...
), the vast majority of them responded to concentration changes in a complex manner. In such instances with several concentration tastants tested, the middle concentration might evoke the highest firing rate (like 0.1 M sucrose), or the highest and lowest concentrations might elicit the highest rates (NaCl ), or the neuron might respond to only one concentration.
The Coherence and Interaction of GC Neurons During Tasting
GC neurons cohere and interact during tasting. GC neurons interact across milliseconds, and these interactions are taste specific and define distinct but overlapping neural assemblies that respond to the presence of each tastant by undergoing coupled changes in firing rate. These couplings are used to discriminate between tastants. Coupled changes in firing rate are the underlying source of GC interactions. Subsets of neurons in GC become coupled after presentation of particular tastants and the responses of neurons in that ensemble change in concert with those of others.Taste familiarity
GC units signal taste familiarity at a delayed temporal phase of the response. An analysis suggests that specific neuronal populations participate in the processing of familiarity for specific tastants. Furthermore, the neural signature of familiarity is correlated with familiarization with a specific tastant rather than with any tastant. This signature is evident 24 hours after initial exposure. This persistent cortical representation of taste familiarity requires slow post-acquisition processing to develop. This process may be related to the activation of neurotransmitterNeurotransmitter
Neurotransmitters are endogenous chemicals that transmit signals from a neuron to a target cell across a synapse. Neurotransmitters are packaged into synaptic vesicles clustered beneath the membrane on the presynaptic side of a synapse, and are released into the synaptic cleft, where they bind to...
receptors, modulation of gene expression, and posttranslational modifications detected in the insular cortex in the first hours after the consumption of an unfamiliar tastant.