Umami Perception Research Articles

EFFECTS OF THREE EMULSION COMPOSITIONS ON TASTE THRESHOLDS AND INTENSITY RATINGS OF FIVE TASTE COMPOUNDS

This study assessed the effects of three emulsified systems on taste thresholds and the near-threshold taste intensities of the five tastes (sweet, sour, salty, bitter and umami). Emulsions were formulated with different lipid chemical compositions. Lipid addition in an emulsified form significantly increased thresholds for sour and bitter tastes produced by citric acid and quinine hydrochloride, respectively. No significant differences were found in the threshold levels in emulsions formulated with different lipids for the five tastes evaluated. In general, for the same tastant concentration, taste intensities for sour and bitter tastes were lower in emulsions compared with the aqueous solutions, which is related to the higher threshold observed in the emulsions. On the other hand, the presence of a lipid phase increased the perception of umami and saltiness, with higher intensity values observed in the emulsions. Finally, lipid addition did not affect sweet intensities. These results suggest that, depending on the type of tastant used, the presence of a lipid phase in an emulsified form may alter the sensory perception of foods. Further research needs to be made for specific products to evaluate the effects of specific ingredients on the sensory properties of foods. PRACTICAL APPLICATIONS Consumers are aware of the health consequences associated with their diets. Current tendencies of incorporating a healthy diet in consumers' everyday lives have challenged food manufacturers to produce foods with improved nutritional qualities while maintaining their sensory characteristics. Some of these improvements include the formulation of low-sodium, low-carbohydrate and/or low-fat products. Understanding the mechanisms associated with taste perception will help formulate new food products with improved sensory and health characteristics.

Pseudogenization of the Umami Taste Receptor Gene Tas1r1 in the Giant Panda Coincided with its Dietary Switch to Bamboo

Although it belongs to the order Carnivora, the giant panda is a vegetarian with 99% of its diet being bamboo. The draft genome sequence of the giant panda shows that its umami taste receptor gene Tas1r1 is a pseudogene, prompting the proposal that the loss of the umami perception explains why the giant panda is herbivorous. To test this hypothesis, we sequenced all six exons of Tas1r1 in another individual of the giant panda and five other carnivores. We found that the open reading frame (ORF) of Tas1r1 is intact in all these carnivores except the giant panda. The rate ratio (ω) of nonsynonymous to synonymous substitutions in Tas1r1 is significantly higher for the giant panda lineage than for other carnivore lineages. Based on the ω change and the observed number of ORF-disrupting substitutions, we estimated that the functional constraint on the giant panda Tas1r1 was relaxed ∼ 4.2 Ma, with its 95% confidence interval between 1.3 and 10 Ma. Our estimate matches the approximate date of the giant panda's dietary switch inferred from fossil records. It is probable that the giant panda's decreased reliance on meat resulted in the dispensability of the umami taste, leading to Tas1r1 pseudogenization, which in turn reinforced its herbivorous life style because of the diminished attraction of returning to meat eating in the absence of Tas1r1. Nonetheless, additional factors are likely involved because herbivores such as cow and horse still retain an intact Tas1r1.

Variation in umami perception and in candidate genes for the umami receptor in mice and humans

The unique taste induced by monosodium glutamate is referred to as umami taste. The umami taste is also elicited by the purine nucleotides inosine 5′-monophosphate and guanosine 5′-monophosphate. There is evidence that a heterodimeric G protein–coupled receptor, which consists of the T1R1 (taste receptor type 1, member 1, Tas1r1) and the T1R3 (taste receptor type 1, member 3, Tas1r3) proteins, functions as an umami taste receptor for rodents and humans. Splice variants of metabotropic glutamate receptors, mGluR1 (glutamate receptor, metabotropic 1, Grm1) and mGluR4 (glutamate receptor, metabotropic 4, Grm4), also have been proposed as taste receptors for glutamate. The taste sensitivity to umami substances varies in inbred mouse strains and in individual humans. However, little is known about the relation of umami taste sensitivity to variations in candidate umami receptor genes in rodents or in humans. In this article, we summarize current knowledge of the diversity of umami perception in mice and humans. Furthermore, we combine previously published data and new information from the single nucleotide polymorphism databases regarding variation in the mouse and human candidate umami receptor genes: mouse Tas1r1 (TAS1R1 for human), mouse Tas1r3 (TAS1R3 for human), mouse Grm1 (GRM1 for human), and mouse Grm4 (GRM4 for human). Finally, we discuss prospective associations between variation of these genes and umami taste perception in both species.

Sensory and receptor responses to umami: an overview of pioneering work

This article provides a selective overview of the early studies of umami taste and outlines significant questions for further research. Umami compounds such as the amino acid glutamate [often in the form of the sodium salt monosodium glutamate (MSG)] and the nucleotide monophosphates 5′-inosinate and 5′-guanylate occur naturally in, and provide flavor for, many foods and cuisines around the world. Early researchers in the United States found that the flavor of pure MSG was difficult to describe. But they all agreed that, although humans found umami compounds, when tasted alone, to be unpalatable, subjects reported that these compounds improved the taste of foods. This taste “dichotomy” may be partly unlearned because it is also observed in very young infants. The uniqueness of umami perception is based on several lines of evidence. First, numerous perceptual studies have shown that the sensation aroused by MSG is distinct from that of the other 4 taste qualities. Second, biochemical studies that show the synergy of the binding of MSG and 5′-guanylate to tongue taste tissue mirror this hallmark perceptual effect. Third, several specific receptors that may mediate umami taste have recently been identified. There remain, however, a number of puzzles surrounding the umami concept, including the molecular basis for an apparent tactile component to umami perception, the reason for the unpalatability of pure umami, and the functional significance for human health and nutrition of umami detection. Future work aimed at understanding these and other open issues will profitably engage scientists in umami research well into the next century.

Taste receptors for umami: the case for multiple receptors

Umami taste is elicited by many small molecules, including amino acids (glutamate and aspartate) and nucleotides (monophosphates of inosinate or guanylate, inosine 5′-monophosphate and guanosine-5′-monophosphate). Mammalian taste buds respond to these diverse compounds via membrane receptors that bind the umami tastants. Over the past 15 y, several receptors have been proposed to underlie umami detection in taste buds. These receptors include 2 glutamate-selective G protein–coupled receptors, mGluR4 and mGluR1, and the taste bud–expressed heterodimer T1R1+T1R3. Each of these receptors is expressed in small numbers of cells in anterior and posterior taste buds. The mGluRs are activated by glutamate and certain analogs but are not reported to be sensitive to nucleotides. In contrast, T1R1+T1R3 is activated by a broad range of amino acids and displays a strongly potentiated response in the presence of nucleotides. Mice in which the Grm4 gene is knocked out show a greatly enhanced preference for umami tastants. Loss of the Tas1r1 or Tas1R3 genes is reported to depress but not eliminate neural and behavioral responses to umami. When intact mammalian taste buds are apically stimulated with umami tastants, their functional responses to umami tastants do not fully resemble the responses of a single proposed umami receptor. Furthermore, the responses to umami tastants persist in the taste cells of T1R3-knockout mice. Thus, umami taste detection may involve multiple receptors expressed in different subsets of taste cells. This receptor diversity may underlie the complex perception of umami, with different mixtures of amino acids, peptides, and nucleotides yielding subtly distinct taste qualities.

Perceptual variation in umami taste and polymorphisms in TAS1R taste receptor genes

BackgroundThe TAS1R1 and TAS1R3 G protein–coupled receptors are believed to function in combination as a heteromeric glutamate taste receptor in humans. ObjectiveWe hypothesized that variations in the umami perception of glutamate would correlate with variations in the sequence of these 2 genes, if they contribute directly to umami taste. DesignIn this study, we first characterized the general sensitivity to glutamate in a sample population of 242 subjects. We performed these experiments by sequencing the coding regions of the genomic TAS1R1 and TAS1R3 genes in a separate set of 87 individuals who were tested repeatedly with monopotassium glutamate (MPG) solutions. Last, we tested the role of the candidate umami taste receptor hTAS1R1-hTAS1R3 in a functional expression assay. ResultsA subset of subjects displays extremes of sensitivity, and a battery of different psychophysical tests validated this observation. Statistical analysis showed that the rare T allele of single nucleotide polymorphism (SNP) R757C in TAS1R3 led to a doubling of umami ratings of 25 mmol MPG/L. Other suggestive SNPs of TAS1R3 include the A allele of A5T and the A allele of R247H, which both resulted in an approximate doubling of umami ratings of 200 mmol MPG/L. We confirmed the potential role of the human TAS1R1-TAS1R3 heteromer receptor in umami taste by recording responses, specifically to l-glutamate and inosine 5′-monophosphate (IMP) mixtures in a heterologous expression assay in HEK (human embryonic kidney) T cells. ConclusionsThere is a reliable and valid variation in human umami taste of l-glutamate. Variations in perception of umami taste correlated with variations in the human TAS1R3 gene. The putative human taste receptor TAS1R1-TAS1R3 responds specifically to l-glutamate mixed with the ribonucleotide IMP. Thus, this receptor likely contributes to human umami taste perception.

Variation in the Human TAS1R Taste Receptor Genes

We have performed a comprehensive evaluation of single-nucleotide polymorphisms (SNPs) and haplotypes in the human TAS1R gene family, which encodes receptors for sweet and umami tastes. Complete DNA sequences of TAS1R1-, TAS1R2-, and TAS1R3-coding regions, obtained from 88 individuals of African, Asian, European, and Native American origin, revealed substantial coding and noncoding diversity: polymorphisms are common in these genes, and polymorphic sites and SNP frequencies vary widely in human populations. The genes TAS1R1 and TAS1R3, which encode proteins that act as a dimer to form the umami (glutamate) taste receptor, showed less variation than the TAS1R2 gene, which acts as a dimer with TAS1R3 to form the sweet taste receptor. The TAS1R3 gene, which encodes a subunit common to both the sweet and umami receptors, was the most conserved. Evolutionary genetic analysis indicates that these variants have come to their current frequencies under natural selection during population growth and support the view that the coding sequence variants affect receptor function. We propose that human populations likely vary little with respect to umami perception, which is controlled by one major form of the receptor that is optimized for detecting glutamate but may vary much more with respect to sweet perception.

Umami: a universal taste1

The identification and acceptance of umami as a basic taste together with the recent advances in umami research are reviewed, beginning with fish sauces in ancient Greece and Rome and concluding with an examination of the data leading to the realization that umami is indeed a basic taste. The contribution of umami to the flavor of a variety of foods, the role of glutamate and 5′-ribonucleotides in triggering the perception of umami as well as the role of glutamate, the dominant umami compound, in human metabolism are also reviewed. Dedicated to the memory of Dr. Roy Teranishi