PMC:1181811 / 5625-19047 JSONTXT

{"target":"https://pubannotation.org/docs/sourcedb/PMC/sourceid/1181811","sourcedb":"PMC","sourceid":"1181811","source_url":"http://www.ncbi.nlm.nih.gov/pmc/1181811","text":"Results\n\nTaste testing\nPrevious efforts to map QTL for bitter taste have utilized consumption tests that may be confounded by the contributions of post-ingestive effects [29]. We used a modified brief-access lick test, which minimizes the contribution of such effects [29,37] to determine whether B6 and D2 mice display differences in taste sensitivity to the taste stimuli QHCl and DB. After initially screening B6 and D2 mice to determine stimulus concentrations that were aversive but not saturating [47], we selected two ligand concentrations for each compound that best differentiated the two strains. Subsequent taste testing of BXD RI lines was restricted to these two concentrations (1 and 3 mM for both QHCl and DB). Avoidance by male and female B6 and D2 mice increased (as indicated by the decreased lick ratio) in a concentration-dependent manner for both compounds (Figure 1A; Table 1). There was a significant strain difference for both 1 and 3 mM QHCl (F[1,25] \u003e 24.6; p \u003c 0.0001). D2 mice displayed decreased aversion relative to B6 mice at both concentrations. On the other hand, the strains did not significantly differ in taste sensitivity to DB (Figure 1A). There were no significant effects of gender.\nTable 1 Mean lick ratios for B6, D2 and BXD mice.\nStrain n Water licks/5s 1 mM DB 3 mM DB 3 mM PR 10 mM PR 1 mM QH 3 mM QH\nB6 16 29.02 ± 1.4 0.606 0.318 0.311 0.164 0.241 0.144\nD2 12 33.91 ± 2.4 0.465 0.286 0.616 0.429 0.758 0.305\nBXD1 5 32.48 ± 1.5 0.372 0.202 0.314 0.252 0.746 0.422\nBXD2 4 36.50 ± 0.9 1.025 0.605 0.208 0.230 0.320 0.215\nBXD5 5 37.23 ± 2.7 0.628 0.354 0.502 0.636 0.836 0.452\nBXD6 5 22.43 ± 3.5 0.442 0.132 0.200 0.124 0.130 0.080\nBXD11 7 34.48 ± 3.9 0.310 0.223 0.387 0.260 0.479 0.260\nBXD13 5 27.74 ± 3.4 0.360 0.290 0.472 0.168 0.692 0.330\nBXD14 7 31.09 ± 3.5 0.741 0.304 0.293 0.174 0.206 0.126\nBXD15 5 37.36 ± 1.5 0.178 0.120 0.468 0.198 0.362 0.222\nBXD20 5 31.26 ± 2.7 0.252 0.200 0.162 0.128 0.150 0.116\nBXD21 6 19.77 ± 1.5 0.390 0.262 0.313 0.133 0.192 0.217\nBXD24 5 33.03 ± 2.4 0.318 0.148 0.490 0.238 0.518 0.370\nBXD27 5 33.46 ± 2.5 0.364 0.434 0.216 0.126 0.146 0.142\nBXD29 3 40.19 ± 1.5 0.257 0.187 0.657 0.173 0.633 0.333\nBXD31 5 19.45 ± 0.8 0.302 0.238 0.152 0.158 0.218 0.194\nBXD32 6 27.22 ± 2.3 0.105 0.107 0.385 0.203 0.405 0.245\nBXD33 6 29.03 ± 2.2 0.757 0.387 0.377 0.237 0.342 0.188\nBXD34 6 23.01 ± 3.4 0.310 0.217 0.843 0.257 0.693 0.355\nThe number of individual mice tested for each strain (n) is listed in the second column. Subsequent columns show the mean lick rate to water during testing (± SEM), mean lick ratio for denatonium benzoate (DB), PROP (6-n-propylthiouracil; PR) and quinine hydrochloride (QH) at each of two concentrations (see Methods for details).\nFigure 1 Lick ratios (mean ± SE) for B6, D2 and BXD strains. (A) Mean lick ratios for B6 (filled circles) and D2 (open circles) mice at two concentrations of QHCl and DB. In all panels, a lower mean lick ratio indicates a greater aversion, and therefore greater taste sensitivity, to the stimulus. For panels B, C, and D, each BXD strain is represented by a different color, and listed in order from least sensitive to most sensitive to 1 mM QHCL. (B) Mean lick ratios for the six BXD strains that are most sensitive to QHCl in this assay. (C) Mean lick ratios for the five BXD strains that are least sensitive to QHCl in this assay. (D) Mean lick ratios for the six BXD strains intermediate in QHCl taste sensitivity to those in (B) and (C). Cutoffs for the three QHCl taster groups were arbitrarily set, as there was a continuity of the phenotype at 1 mM QHCl: sensitive strains exhibited a lick ratio for 1 mM QHCl of ≤ 0.3, intermediate strains from 0.31-0.6, and insensitive strains \u003e 0.6. The absence of two distinct phenotypic classes suggests that sensitivity to QHCL is under polygenic control. We next tested mice from 17 BXD lines in the same manner. BXD mice also typically avoided both stimuli in a concentration dependent manner (Figures 1B–1D; Table 1). However, QHCl and DB taste sensitivity vary independently across these RI strains: some strains highly sensitive to QHCl are relatively insensitive to DB, and vice versa (Figures 1B–1D).\n\nQTL mapping\nLinkage analysis was conducted using Map Manager QTX (version 0.30[38]). No significant QTLs were identified for DB taste sensitivity, although several associations with markers on chromosomes 2,8 and 12 were \"suggestive\" (LRS \u003e 9.4, genome-wide p = 0.65; see Additional File 1). A significant (LRS \u003e 20.5; genome-wide p = 0.05) QTL for sensitivity to 1 mM QHCl was indicated on chromosome 6, with a second, suggestive (LRS \u003e 11.4; genome-wide p = 0.65) QTL on chromosome 8 (Figure 2A); at 3 mM QHCl, both of these QTL were suggestive (LRS \u003e 10.9) but did not reach genome-wide significance (Figure 2B).\nFigure 2 A major QTL for QHCl taste on mouse chromosome 6. (Top panel) The interval map (see Methods) shows a significant QTL on chromosome 6 (green) and a suggestive QTL on chromosome 8 (yellow) affecting taste responses to 1 mM QHCl. (Bottom panel) For 3 mM QHCl, both QTL were suggestive (yellow). The dashed line indicates genome-wide significance. The chromosome 6 QTL was linked to a single marker, D6Mit13 (Table 2, Figure 3). Adjacent proximal markers D6Mit254 and D6Mit194 are unlinked to the QHCl QTL, as is distal marker D6Mit374. Across the 17 RI lines tested there is at least one recombination event between D6Mit13 and either D6Mit254 (and D6Mit194, the physical position of which is not well defined) or D6Mit374. An additional proximal marker, D6Mit61, which lies between D6Mit194 and D6Mit13, was identified from genotypes of the BXD lines reported by the Jackson Laboratories. BXD/Ty-34 RI mice display a clear D2 phenotype for QHCl taste (Figure 1C) and D2 genotype for D6Mit13, but have a B6 genotype for D6Mit61 [39,40], indicating that D6Mit61 is unlinked to the QHCl QTL. Therefore, this QTL interval can be conservatively defined as that portion of mouse chromosome 6 that lies between D6Mit254 and D6Mit374, but is most likely restricted to the region between D6Mit61 and D6Mit374.\nPhysical mapping of the single linked marker, D6Mit13, and the two closest unlinked markers, D6Mit61and D6Mit374, was performed in silico based on the May, 2004 build of the public B6 genome. Based on these marker positions, the size of the QHCl chromosome 6 QTL is less than 5.0 Mb (Figure 3). This region contains a number of known genes, all but eleven of which encode members of two large receptor families: natural killer cell lectin-like receptors, and T2R-type taste receptors. The Tas2r genes (which encode the T2Rs) are found clustered within a 1.2 Mb interval on either side of D6Mit13 (Figure 3, Figure 4). Because of their proximity to the linked marker, their demonstrated expression in taste receptor cells, and their role in the detection of at least some bitter-tasting compounds, we hypothesized that one or more of the 24 Tas2rs at this locus were responsible for the major QHCl taste sensitivity QTL.\nFigure 3 The QHCl QTL is linked to a single marker on chromosome 6. (A) As shown in the interval map for chromosome 6, the trait value (lick ratio for 1 mM QHCl) correlates strongly across BXD RI strains with the polymorphic marker D6Mit13 (bold). The dashed line indicates genome-wide significance. (B) The QHCl QTL (which lies between unlinked markers D6Mit61 and D6Mit 374) contains a cluster of putative bitter taste receptor genes, the Tas2rs (gray box). Physical positions of the polymorphic markers are given in Mb, and are based on the May, 2004 build of the B6 mouse genome. The physical position of D6Mit194 (*) is tentative.\nFigure 4 A map of the distal chromosome 6 Tas2r cluster. Twenty-four intact Tas2r genes map to distal chromosome 6 (black). The Tas2rs are found in two subclusters on either side of the polymorphic marker D6Mit13 (red) and two genes encoding proline-rich salivary proteins (Prp2 and Prh1; red). Map positions, in Mb, represent chromosome 6 positions in the May, 2004 assembly of the B6 genome.\n\nT2R alleles\nIf one (or more) Tas2rs underlie the chromosome 6 QHCl taste sensitivity QTL, we would predict that one (or more) Tas2r genes would exhibit one of three likely characteristics: (1) A Tas2r allele is a pseudogene, or is deleted, in D2 (QHCL-insensitive), but not B6 (QHCl-sensitive), mice; (2) Missense mutations in the single coding exon of a D2 Tas2r allele impact protein functions such as ligand binding or receptor coupling to downstream signaling cascades; (3) Mutations in noncoding or regulatory regions of a D2 Tas2r allele affects expression of the protein product. Though we considered all three of these to be valid possibilities, we initially focused on the likelihood that deletion or mutation within the coding sequence of a single D2 Tas2r would correlate with the QHCl taste insensitivity phenotype.\nTwenty-four intact Tas2rs, along with three apparent Tas2r pseudogenes, have been identified in the distal chromosome 6 cluster of B6 mice [19] (Figure 4). We designed oligonucleotides to non-coding regions flanking the coding sequence of each intact Tas2r [see Additional file 2]. Using these oligos, we amplified each Tas2r coding sequence from D2 genomic DNA. PCR products were subcloned into cloning vectors and sequenced. Comparisons of the sequences of B6 and D2 orthologues revealed that only two of the twenty-four Tas2r alleles examined, Tas2r106 and Tas2r124, were identical across strains at the amino acid level (data not shown). A third, Tas2r120, could not be amplified from D2 genomic DNA (Figure 5) using either of two pairs of oligonucleotides (Additional file 2), suggesting that this Tas2r is deleted in D2 mice. Two D2 alleles, Tas2r103 and Tas2r117, contained numerous missense mutations and small deletions that create frame shifts and premature termination; these two genes may be pseudogenes in this strain. The remaining 19 Tas2rs contained between one and 16 missense mutations. All 24 Tas2rs examined have different alleles in B6 and D2 mice, and 307 single nucleotide polymorphisms are present within coding exons (data not shown). Although polymorphic residues between B6 and D2 Tas2rs are found in all regions of the receptors, 23% of the amino acid changes seen are within the first two extracellular loops of the T2Rs (data not shown).\nFigure 5 Allelic variation across strains for four Tas2rs. B6 and D2 alleles of four Tas2rs can be differentiated based on diagnostic restriction digests of amplified PCR products (Tas2r105, Tas2r116 and Tas2r131) or on the presence or absence of a PCR product (Tas2r120). In each of the 17 BXD strains tested, Tas2r genotype was always correlated with QHCl taster phenotype (blue = B6 taster phenotype, red = D2 taster phenotype). See additional file 1: Table 3 for restriction enzymes and oligonucleotides. The variability between orthologous receptors in these two inbred strains suggested that it might be possible to narrow the physical boundaries of the QHCl taste QTL by determining which Tas2r alleles are correlated with QHCl taste sensitivity. Therefore, we proceeded to screen genomic DNA from 29 available BXD RI lines, including the 17 that we had used in taste testing, for the Tas2r alleles they contained. In most cases, we were able to identify diagnostic restriction endonuclease digests that would allow us to quickly identify whether a particular Tas2r PCR product was amplified from a B6 or D2 allele. We did not analyze three genes (Tas2r104, Tas2r114 and Tas2r110) where no diagnostic restriction endonuclease could be identified. For Tas2r120, which is likely deleted in D2 mice, the absence of a PCR product was diagnostic of the D2 genotype for this gene.\nSurprisingly, we discovered that there have been no apparent recombination events within the distal chromosome 6 cluster during the generation of the BXD RI lines. For all RI lines tested, every Tas2r within an individual RI line originated from the same parental strain (Figures 5, 6). Furthermore, the genotype of each Tas2r gene always correlated with the QHCl taste phenotype (Figures 6, 7), suggesting that the entire Tas2r cluster is a single haplotype that varies with QHCl taster status.\nFigure 6 The Tas2r cluster is a single haplotype in BXD/Ty RI mice. The coding exon of each of 21 Tas2rs in the distal chromosome 6 cluster was amplified genomic DNA from 29 BXD/Ty RI strains. Each Tas2r within an individual BXD strain originated from the same parental strain (B6 allele = gray, D2 allele = white). The 17 BXD strains that were behaviorally tested in this study are indicated (*).\nFigure 7 Tas2r genotype correlates with QHCl taste phenotype. Mean lick ratios of B6, D2 and BXD strains reported in Figure 1 are grouped based on Tas2r haplotype (B6 haplotype = blue, D2 haplotype = red). B6 mice (blue line on left panel) are more sensitive to 1 mM and 3 mM QHCl than are D2 mice (red line on left panel) in brief access taste tests. Similarly, BXD strains with the B6 Tas2r haplotype (blue lines, right panel) are more sensitive to QHCl than are BXD strains with the D2 Tas2r haplotype (red lines, right panel). The BXD strains are listed in order from least to most sensitive to 1 mM 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