Biochemistry of Gustation

Taste
• common language “taste” describe sensations arising
from the oral cavity
• biological definition of taste, or gustation, is narrower and
includes:
– only sensations mediated by a specialized anatomically and
physiologically defined chemosensory gustatory system
 Taste
• along with taste sensations:
• food usually simultaneously evokes other sensations:
– Odor
– Touch
– Temperature
– Irritation
• it’s not always easy to separate all these sensations
perceptually
• the nongustatory components are sensed by different
systems:
– olfaction
– somatosensation
• taste and olfaction:
– among the five senses:
• play crucial roles in the detection of chemical
substances in the environment:
– are referred to as chemical senses
circumvallate papilla
 Labelled Lines
• 5 recognised modalities:
– sweet, sour, bitter, salt & umami
– all mature taste bud cells have prominent microvilli
• Taste bud cells:
– are continuously renewed from a local population of
stem cells
– have half-life of about ten days
• Type I "dark" cells:
– are glial (supporting) cells
– also contribute to salty (chloride) tastes
– full salt flavour may also require sodium channels
 Labelled Lines
• Sub-populations of type II cells:
– respond to sweet, bitter, umami (glutamate) and
possibly fats
– they release ATP which stimulates purinergic receptors
on:
• type III cells
• also on sensory neurons

• Type III cells:

– signal "sour" after intracellular acidification
– only type III cells make synaptic connections where they
release serotonin
 Labelled Lines
• sensory nerves:
– are “labelled lines”
– but it is not known how they specifically communicate
with the appropriate sensory cells
Taste bud
 Recognised tastes & receptors
• Sour – PKD1L3 or PKD2L1 (disputed)
• Salty – ENaC or TRPV1 (disputed)
• Bitter – T2R family – about 30 members
• Sweet – T1R2/T1R3 heterodimer
• Umami – T1R1/T1R3 heterodimer
– plus other glutamate receptors?

• Fats?

• In addition to the taste buds, there is a closely-related

family of entero-endocrine cells throughout the GI tract,
which use similar transduction mechanisms to control
food intake, digestion and incretin release.
 Human Taste Sensations
• divided into five qualities:
– Bitter:
• aversive bitter taste often indicates presence of toxins in food
– Sour:
• Together with sour taste, may also signal spoiled food
– Salty:
• main salty taste stimuli are sodium salts, but some nonsodium salts also
have a salty taste component:
– suggests that salty taste signals the presence of either sodium or minerals in general

– Sweet:
• most common natural sweet taste stimuli are sugars, which indicate the
presence of carbohydrates in food
– Umami:
• (savory; the prototypical stimulus being the amino acid glutamate)
 Discovery of Taste Receptors
• Fuller:
– in 1974, using long-term two-bottle tests

• A positional cloning study at the Monell

Chemical Senses Center
– second line of studies stemmed from analyses of a
taste-bud-enriched cDNA library (72), which resulted
in a discovery of two putative G protein–coupled taste
receptors, T1R1 and T1R2
Nomenclature & Classification of Taste
Receptor Genes & Proteins
• two best-characterized families of taste
receptors:
• T1R - “taste receptor, type 1”
• T2R - “taste receptor, type 2”
• Tas1r1(in mouse or rat) or TAS1R1 (in human):
– standard gene names follow the description: “taste
receptor, type 1” (with corresponding type and
member numbers).
• T1R1:
– a corresponding protein symbol is (uppercase letters
and not italicized)
 Classification Systems
• Several classification systems for the GPCR
superfamily:
– one of the most frequently used includes GPCRs:
• six classes (clans): A, B, C, D, E, and F
– according to this classification:
• T1Rs belong to class C (metabotropic glutamate/pheromone)
GPCRs
• T2Rs are described either as a separate putative family or as
distantly related to class A (rhodopsinlike) GPCRs
 Classification Systems
• GRAFTS (glutamaterhodopsin-adhesion-
frizzled/taste2-secretin) classification system:
– more recent
– developed based on phylogenetic analyses of
transmembrane parts of human GPCRs
– according to this classification:
• T1Rs belong to the glutamate family
• T2Rs form a distinct cluster within the frizzled/taste2 family
– the second cluster of this family includes:
• frizzled receptors involved in cell proliferation and
development
Bitter, sweet & umami taste transduction
26
 Gustatory Adaptation
• It is difficult to study sensory adaptation to continuing taste stimulation
and there are very few published reports.
• It has recently become apparent that taste sensitivity is modulated by
external factors, and this has become a prolific area for research.
• Leptin, a cytokine produced by adipocytes (which reduces food intake
and increases energy expenditure) inhibits the taste bud response to
saccharin.
• Glucagon (which is produced in response to hypoglycaemia) increases
the taste bud response to sucrose, while the incretin hormone GLP-1
maintains and enhances the response to sweet stimuli.
• Ghrelin increases the response to salty and sour tastants, and oxytocin
also seems likely to modulate these responses.
• Taste buds also respond to GABA, and it seems likely that complex
networks of signalling molecules will be found to underpin taste bud
physiology, just as they underpin other areas of the GI tract.