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gustatory system of mammals is thought to detect thousands of chemically-diverse bitter-tasting substances [41]. Although specific receptors, enzymes and channels have been implicated in the transduction of bitter stimuli, how interactions of bitter stimuli with taste receptor cells lead to cellular activation and signaling to the central nervous system is still poorly understood. We have found that a single QTL on distal chromosome 6 accounts for most of the variation in QHCL taste sensitivity between B6 and D2 mice. This QTL maps to the same chromosomal position as a previously identified QTL for quinine intake, Qui [16,28], indicating that taste is the major factor in regulating quinine aversion. This is an important distinction, as the consumption of bitter-tasting stimuli can be dependent on factors independent of taste, such as toxicity [29].\nUsing 17 RI lines and 762 chromosomal markers, we have restricted the quinine taste QTL to a \u003c 5 Mb region on distal chromosome 6 that contains 24 Tas2r genes. At least 60 other genes also lie within this interval, including two genes that encode proline-rich salivary proteins, Prp2 and Prh1; these proteins appear to play no direct role in bitter taste [48]. Tas2rs are the most likely candidates for the QHCl quantitative trait gene(s) due to: (1) their expression in taste receptor cells and (2) genetic and functional evidence linking them to the detection of a number of bitter taste stimuli. As of yet there is no evidence for quinine activation of T2Rs from functional assays of these receptors in heterologous cells or membrane preparations, likely due to the lipophilic nature of quinine [23]. However, several physiological studies have suggested that quinine can directly activate G proteins or cationic conductances, or can block K+ channels in taste receptor cells [34-36]. While our data indicates that quinine taste is largely T2R-dependent, it is not exclusively so. For example, the BXD RI lines exhibited a range of quinine sensitivity, with several strains having similar sensitivities to that of B6, some strains with sensitivities similar to that of D2, and a third group with a more intermediate phenotype (Figures 1, 7). This observation is consistent with a polygenic basis for quinine taste [16,26]. Also, a suggestive QTL on chromosome 8 does not contain any Tas2r genes, but does contain a number of genes encoding ion channels, enzymes and members of other receptor families (our unpublished data). It will be interesting to determine whether this suggestive QTL is linked to quinine taste and, if so, whether it is specific for this single bitter stimulus or more broadly related to all bitter taste.\nOf the 29 BXD RI lines examined, there was no apparent recombination event within the chromosome 6 Tas2r cluster. While increasing the number of BXD RI lines or the number of markers used for genotyping them would facilitate the definition of smaller QTL intervals, in this case such an effort is unlikely to permit the identification of one or a few Tas2rs involved in quinine taste. For example, we examined six lines of AXB and BXA RIs with reported recombinations around D6Mit13; a small sampling of the Tas2rs in these RI lines again indicated no recombinations within the Tas2r cluster (data not shown). Behavioral genetic approaches have been invaluable for identifying genes involved in taste function, such as the Tas1r3 gene that encodes a receptor important for sweet and umami taste [42]. Positional cloning also permitted the identification of the Tas2r responsible for the majority of variance of phenylthiocarbamide (PTC) taste sensitivity in humans [18]. In both of these cases, however, the genes linked to saccharin or PTC taste were not tightly clustered with paralogues. For bitter taste, behavioral genetic approaches may be more useful for identifying genes encoding downstream signaling molecules or components of T2R-independent transduction mechanisms. For example, a QTL for PROP avoidance has been suggested on chromosome 7 [16], and we observe a suggestive QTL for quinine taste on chromosome 8 (Figure 2); in neither case are Tas2rs found at these loci (data not shown).\nIt is somewhat puzzling that 22 of the 24 Tas2rs examined encode variant proteins in B6 and D2 mice even though these strains exhibit similar taste responses to bitter compounds such as DB or cyclohexamide [47]. Taken together, these observations suggest that Tas2rs are quite tolerant of variation, and that perhaps most of the differences observed do not affect domains important for ligand interactions or receptor-mediated signaling mechanisms. Interestingly, 23% of missense mutations in D2 Tas2rs affect the first two extracellular loops of the receptors. These two loops have been recently shown to affect the ligand response profiles of some T2Rs [23]. More systematic analyses of structure-function relationships between these T2R variants and an array of bitter stimuli are necessary to determine which changes may impact ligand binding, interactions with other proteins, or overall receptor structure.\nSuch large numbers of nonsynonymous substitutions between orthologues is suggestive of adaptive selection. Analysis of sequence diversity of Tas2rs in humans, great apes and old world monkeys suggest that Tas2rs are subject to some degree of positive selection [43,44]. However, the fact that these two mouse strains, members of the same species, are so closely related makes this explanation problematic. It is possible that B6 and D2 mice, which have a similar origin in the early 20th century, inherited different Tas2r haplotypes present in wild mouse populations prior to inbreeding. Characterization of Tas2r sequences of several wild mouse species or subspecies, or in other inbred lines, would shed light on this issue."}

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gustatory system of mammals is thought to detect thousands of chemically-diverse bitter-tasting substances [41]. Although specific receptors, enzymes and channels have been implicated in the transduction of bitter stimuli, how interactions of bitter stimuli with taste receptor cells lead to cellular activation and signaling to the central nervous system is still poorly understood. We have found that a single QTL on distal chromosome 6 accounts for most of the variation in QHCL taste sensitivity between B6 and D2 mice. This QTL maps to the same chromosomal position as a previously identified QTL for quinine intake, Qui [16,28], indicating that taste is the major factor in regulating quinine aversion. This is an important distinction, as the consumption of bitter-tasting stimuli can be dependent on factors independent of taste, such as toxicity [29].\nUsing 17 RI lines and 762 chromosomal markers, we have restricted the quinine taste QTL to a \u003c 5 Mb region on distal chromosome 6 that contains 24 Tas2r genes. At least 60 other genes also lie within this interval, including two genes that encode proline-rich salivary proteins, Prp2 and Prh1; these proteins appear to play no direct role in bitter taste [48]. Tas2rs are the most likely candidates for the QHCl quantitative trait gene(s) due to: (1) their expression in taste receptor cells and (2) genetic and functional evidence linking them to the detection of a number of bitter taste stimuli. As of yet there is no evidence for quinine activation of T2Rs from functional assays of these receptors in heterologous cells or membrane preparations, likely due to the lipophilic nature of quinine [23]. However, several physiological studies have suggested that quinine can directly activate G proteins or cationic conductances, or can block K+ channels in taste receptor cells [34-36]. While our data indicates that quinine taste is largely T2R-dependent, it is not exclusively so. For example, the BXD RI lines exhibited a range of quinine sensitivity, with several strains having similar sensitivities to that of B6, some strains with sensitivities similar to that of D2, and a third group with a more intermediate phenotype (Figures 1, 7). This observation is consistent with a polygenic basis for quinine taste [16,26]. Also, a suggestive QTL on chromosome 8 does not contain any Tas2r genes, but does contain a number of genes encoding ion channels, enzymes and members of other receptor families (our unpublished data). It will be interesting to determine whether this suggestive QTL is linked to quinine taste and, if so, whether it is specific for this single bitter stimulus or more broadly related to all bitter taste.\nOf the 29 BXD RI lines examined, there was no apparent recombination event within the chromosome 6 Tas2r cluster. While increasing the number of BXD RI lines or the number of markers used for genotyping them would facilitate the definition of smaller QTL intervals, in this case such an effort is unlikely to permit the identification of one or a few Tas2rs involved in quinine taste. For example, we examined six lines of AXB and BXA RIs with reported recombinations around D6Mit13; a small sampling of the Tas2rs in these RI lines again indicated no recombinations within the Tas2r cluster (data not shown). Behavioral genetic approaches have been invaluable for identifying genes involved in taste function, such as the Tas1r3 gene that encodes a receptor important for sweet and umami taste [42]. Positional cloning also permitted the identification of the Tas2r responsible for the majority of variance of phenylthiocarbamide (PTC) taste sensitivity in humans [18]. In both of these cases, however, the genes linked to saccharin or PTC taste were not tightly clustered with paralogues. For bitter taste, behavioral genetic approaches may be more useful for identifying genes encoding downstream signaling molecules or components of T2R-independent transduction mechanisms. For example, a QTL for PROP avoidance has been suggested on chromosome 7 [16], and we observe a suggestive QTL for quinine taste on chromosome 8 (Figure 2); in neither case are Tas2rs found at these loci (data not shown).\nIt is somewhat puzzling that 22 of the 24 Tas2rs examined encode variant proteins in B6 and D2 mice even though these strains exhibit similar taste responses to bitter compounds such as DB or cyclohexamide [47]. Taken together, these observations suggest that Tas2rs are quite tolerant of variation, and that perhaps most of the differences observed do not affect domains important for ligand interactions or receptor-mediated signaling mechanisms. Interestingly, 23% of missense mutations in D2 Tas2rs affect the first two extracellular loops of the receptors. These two loops have been recently shown to affect the ligand response profiles of some T2Rs [23]. More systematic analyses of structure-function relationships between these T2R variants and an array of bitter stimuli are necessary to determine which changes may impact ligand binding, interactions with other proteins, or overall receptor structure.\nSuch large numbers of nonsynonymous substitutions between orthologues is suggestive of adaptive selection. Analysis of sequence diversity of Tas2rs in humans, great apes and old world monkeys suggest that Tas2rs are subject to some degree of positive selection [43,44]. However, the fact that these two mouse strains, members of the same species, are so closely related makes this explanation problematic. It is possible that B6 and D2 mice, which have a similar origin in the early 20th century, inherited different Tas2r haplotypes present in wild mouse populations prior to inbreeding. Characterization of Tas2r sequences of several wild mouse species or subspecies, or in other inbred lines, would shed light on this issue."}

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gustatory system of mammals is thought to detect thousands of chemically-diverse bitter-tasting substances [41]. Although specific receptors, enzymes and channels have been implicated in the transduction of bitter stimuli, how interactions of bitter stimuli with taste receptor cells lead to cellular activation and signaling to the central nervous system is still poorly understood. We have found that a single QTL on distal chromosome 6 accounts for most of the variation in QHCL taste sensitivity between B6 and D2 mice. This QTL maps to the same chromosomal position as a previously identified QTL for quinine intake, Qui [16,28], indicating that taste is the major factor in regulating quinine aversion. This is an important distinction, as the consumption of bitter-tasting stimuli can be dependent on factors independent of taste, such as toxicity [29].\nUsing 17 RI lines and 762 chromosomal markers, we have restricted the quinine taste QTL to a \u003c 5 Mb region on distal chromosome 6 that contains 24 Tas2r genes. At least 60 other genes also lie within this interval, including two genes that encode proline-rich salivary proteins, Prp2 and Prh1; these proteins appear to play no direct role in bitter taste [48]. Tas2rs are the most likely candidates for the QHCl quantitative trait gene(s) due to: (1) their expression in taste receptor cells and (2) genetic and functional evidence linking them to the detection of a number of bitter taste stimuli. As of yet there is no evidence for quinine activation of T2Rs from functional assays of these receptors in heterologous cells or membrane preparations, likely due to the lipophilic nature of quinine [23]. However, several physiological studies have suggested that quinine can directly activate G proteins or cationic conductances, or can block K+ channels in taste receptor cells [34-36]. While our data indicates that quinine taste is largely T2R-dependent, it is not exclusively so. For example, the BXD RI lines exhibited a range of quinine sensitivity, with several strains having similar sensitivities to that of B6, some strains with sensitivities similar to that of D2, and a third group with a more intermediate phenotype (Figures 1, 7). This observation is consistent with a polygenic basis for quinine taste [16,26]. Also, a suggestive QTL on chromosome 8 does not contain any Tas2r genes, but does contain a number of genes encoding ion channels, enzymes and members of other receptor families (our unpublished data). It will be interesting to determine whether this suggestive QTL is linked to quinine taste and, if so, whether it is specific for this single bitter stimulus or more broadly related to all bitter taste.\nOf the 29 BXD RI lines examined, there was no apparent recombination event within the chromosome 6 Tas2r cluster. While increasing the number of BXD RI lines or the number of markers used for genotyping them would facilitate the definition of smaller QTL intervals, in this case such an effort is unlikely to permit the identification of one or a few Tas2rs involved in quinine taste. For example, we examined six lines of AXB and BXA RIs with reported recombinations around D6Mit13; a small sampling of the Tas2rs in these RI lines again indicated no recombinations within the Tas2r cluster (data not shown). Behavioral genetic approaches have been invaluable for identifying genes involved in taste function, such as the Tas1r3 gene that encodes a receptor important for sweet and umami taste [42]. Positional cloning also permitted the identification of the Tas2r responsible for the majority of variance of phenylthiocarbamide (PTC) taste sensitivity in humans [18]. In both of these cases, however, the genes linked to saccharin or PTC taste were not tightly clustered with paralogues. For bitter taste, behavioral genetic approaches may be more useful for identifying genes encoding downstream signaling molecules or components of T2R-independent transduction mechanisms. For example, a QTL for PROP avoidance has been suggested on chromosome 7 [16], and we observe a suggestive QTL for quinine taste on chromosome 8 (Figure 2); in neither case are Tas2rs found at these loci (data not shown).\nIt is somewhat puzzling that 22 of the 24 Tas2rs examined encode variant proteins in B6 and D2 mice even though these strains exhibit similar taste responses to bitter compounds such as DB or cyclohexamide [47]. Taken together, these observations suggest that Tas2rs are quite tolerant of variation, and that perhaps most of the differences observed do not affect domains important for ligand interactions or receptor-mediated signaling mechanisms. Interestingly, 23% of missense mutations in D2 Tas2rs affect the first two extracellular loops of the receptors. These two loops have been recently shown to affect the ligand response profiles of some T2Rs [23]. More systematic analyses of structure-function relationships between these T2R variants and an array of bitter stimuli are necessary to determine which changes may impact ligand binding, interactions with other proteins, or overall receptor structure.\nSuch large numbers of nonsynonymous substitutions between orthologues is suggestive of adaptive selection. Analysis of sequence diversity of Tas2rs in humans, great apes and old world monkeys suggest that Tas2rs are subject to some degree of positive selection [43,44]. However, the fact that these two mouse strains, members of the same species, are so closely related makes this explanation problematic. It is possible that B6 and D2 mice, which have a similar origin in the early 20th century, inherited different Tas2r haplotypes present in wild mouse populations prior to inbreeding. Characterization of Tas2r sequences of several wild mouse species or subspecies, or in other inbred lines, would shed light on this issue."}