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Research ArticleNew Research, Cognition and Behavior

Nonphosphorylatable Src Ser75 Mutation Increases Ethanol Preference and Consumption in Mice

Goro Kato
eNeuro 28 March 2019, 6 (2) ENEURO.0418-18.2019; https://doi.org/10.1523/ENEURO.0418-18.2019
Goro Kato
Department of Biochemistry, Faculty of Medicine, Graduate Faculty of Interdisciplinary Research, University of Yamanashi, Yamanashi 409-3898, Japan
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    Figure 1.

    SA/SA mutant mice show increased ethanol consumption. Volitional ethanol intake was measured in a two-bottle choice test between tap water and 5% or 10% ethanol for three weeks in Src Ser75 mutant mice. A, Consumption of a 5% or 10% ethanol solution by SA/SA (left) and SD/SD (right) mice and their WT/WT counterparts. Measurements for mutants were normalized to the average amount per kilogram body weight per measurement period (3 or 4 d) for the WT littermate controls. B, Ethanol preference ratios (volume of ethanol solution consumed per total volume of fluid consumed) of a 5% or 10% ethanol solution in SA/SA (left) and SD/SD (right) mice and their WT/WT counterparts. C, Total fluid intake of a 5% or 10% ethanol solution and water for SA/SA (left) and SD/SD (right) mice and their WT/WT counterparts. Measurements for mutants were normalized to the average amount per kilogram body weight per measurement period for WT littermate controls. Black bars, WT/WT; white bars, SA/SA or SD/SD. Data are mean ± SD. Sample numbers are shown in parentheses; *p < 0.05, **p < 0.01 versus WT/WT by unpaired two-tailed Welch’s t tests, except for consumption of 10% ethanol in SD/SD mice, which was analyzed by Mann–Whitney U test.

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    Figure 2.

    Taste preference ratios for sucrose and quinine in WT and SA/SA (A) and SD/SD (B) mutant mice. Taste preference was assessed via a two-bottle choice paradigm with tap water and a tastant, each for 14 d, in the following order: 0.033% (w/v) sucrose, 0.1% (w/v) sucrose, 0.02 mM quinine, and 0.04 mM quinine. C, Total intake of a 0.033% (w/v) or 0.1% (w/v) sucrose solution in SA/SA (left) and SD/SD (right) mice and their WT/WT counterparts. Measurements for mutants were normalized to the average amount of 0.033% (w/v) sucrose per kilogram body weight per measurement period for WT littermate controls. Data are mean ± SD analyzed by paired factorial ANOVA and Bonferroni’s post hoc tests.

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    Figure 3.

    Plasma ethanol levels assayed 1 and 3 h after WT/WT and SA/SA (left) or SD/SD (right) mice were injected with ethanol (3 g/kg). Blood samples (10 μl) were collected from each mouse at the postinjection time points and used for the alcohol dehydrogenase enzymatic spectrophotometric assay. Data are mean ± SD analyzed by paired factorial ANOVA and Tukey–Kramer post hoc tests.

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    Figure 4.

    LORR for SA/SA (left) and SD/SD (right) mice and their WT counterparts after they were injected with ethanol (3 g/kg). Data are mean ± SD analyzed by unpaired and two-tailed Welch’s t tests.

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    Figure 5.

    Downregulation of ROCK signaling in striatal tissues from SA/SA mutant mice. A, ROCK activity levels in the striatal tissues of SA/SA (left) and SD/SD (right) mutant mice measured by immunoassays. B, ROCK protein levels in striatal tissues from SA/SA (left) and SD/SD (right) mutant mice measured by Western blotting (upper). ROCK protein level (A.U.) in each tissue was normalized to that of β-tubulin (Tub, lower). C, ROCK kinase activity (mU) per milligram of striatal tissues and normalized to ROCK protein amount (bottom) in SA/SA (left) and SD/SD (right) mutant mice. Scatter diagrams of the two variables are shown in upper panels. D, Phosphorylation of 60-kDa Akt at Ser473 (P-Akt) in the striatal tissues of SA/SA mutant mice as measured by Western blotting. Akt phosphorylation levels were normalized to the level of total Akt. Data are mean ± SD; *p < 0.05, **p < 0.01 by unpaired two-tailed Welch’s t tests.

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    Table 1.

    Statistical table

    Figure/tableData structureType of testSample sizeStatistical data
    Fig. 1A
    Ethanol consumption
    WT and SA/SA mice (5%)Normal distributionWelch t test unpairedWT: n = 6, SA/SA: n = 7CI(d): –2.681 and –0.273, t = –2.867, p = 0.0226
    WT and SA/SA mice (10%)Normal distributionWelch t test unpairedWT: n = 5, SA/SA: n = 5CI(d): –1.730 and –0.258, t = –3.562, p = 0.0184
    WT and SD/SD mice (5%)Normal distributionWelch t test unpairedWT: n = 7, SD/SD: n = 6CI(d): –1.705 and 0.765, t = –0.906, p = 0.3960
    WT and SD/SD mice (10%)Non-normal distributionMann–Whitney U testWT: n = 8, SD/SD: n = 8U = 26, p = 0.5286
    Fig. 1B
    Ethanol preference ratio
    WT and SA/SA mice (5%)Normal distributionWelch t test unpairedWT: n = 6, SA/SA: n = 7CI(d): –0.444 and –0.056, t = –2.987, p = 0.0179
    WT and SA/SA mice (10%)Normal distributionWelch t test unpairedWT: n = 5, SA/SA: n = 5CI(d): –0.177 and –0.039, t = –3.724, p = 0.0080
    WT and SD/SD mice (5%)Normal distributionWelch t test unpairedWT: n = 7, SD/SD: n = 6CI(d): –0.311 and 0.187, t = –0.588, p = 0.5747
    WT and SD/SD mice (10%)Normal distributionWelch t test unpairedWT: n = 8, SD/SD: n = 8CI(d): –0.109 and 0.094, t = –0.155, p = 0.8791
    Fig. 1C
    Total liquid intake
    WT and SA/SA mice (5%)Normal distributionWelch t test unpairedWT: n = 6, SA/SA: n = 7CI(d): –0.130 and 0.296, t = 0.916, p = 0.3900
    WT and SA/SA mice (10%)Normal distributionWelch t test unpairedWT: n = 5, SA/SA: n = 5CI(d): –0.156 and 0.133, t = –0.190, p = 0.8546
    WT and SD/SD mice (5%)Normal distributionWelch t test unpairedWT: n = 7, SD/SD: n = 6CI(d): –0.272 and 0.117, t = –0.877, p = 0.3999
    WT and SD/SD mice (10%)Normal distributionWelch t test unpairedWT: n = 8, SD/SD: n = 8CI(d): –0.223 and 0.191, t = –0.166, p = 0.8712
    Fig. 2A
    Taste preference ratio (WT and SA/SA)
    SucroseNormal distributionPaired factorial ANOVAWT: n = 5, SA/SA: n = 5Genotype: F = 0.5107, p = 0.4951
    0.033% (w/v)Bonferroni’s post hocCI(m): WT, 0.477 and 0.569; SA/SA, 0.489 and 0.580, p = 0.5901
    0.1% (w/v)Bonferroni’s post hocCI(m): WT, 0.430 and 0.521; SA/SA, 0.447 and 0.538, p = 0.4224
    QuinineNormal distributionPaired factorial ANOVAWT: n = 5, SA/SA: n = 5Genotype: F = 0.0331, p = 0.8602
    0.02 mMBonferroni’s post hocCI(m): WT, 0.134 and 0.522; SA/SA, 0.139 and 0.527, p = 0.9484
    0.04 mMBonferroni’s post hocCI(m): WT, 0.033 and 0.421; SA/SA, 0.060 and 0.448, p = 0.6783
    Fig. 2B
    Taste preference ratio (WT and SD/SD)
    SucroseNormal distributionPaired factorial ANOVAWT: n = 5, SD/SD: n = 5Genotype: F = 0.8475, p = 0.3842
    0.033% (w/v)Bonferroni’s post hocCI(m): WT, 0.495 and 0.604; SD/SD, 0.447 and 0.557, p = 0.0533
    0.1% (w/v)Bonferroni’s post hocCI(m): WT, 0.518 and 0.627; SD/SD, 0.522 and 0.631, p = 0.8722
    QuinineNormal distributionPaired factorial ANOVAWT: n = 5, SD/SD: n = 5Genotype: F = 0.4920, p = 0.5029
    0.02 mMBonferroni’s post hocCI(m): WT, 0.231 and 0.540; SD/SD, 0.190 and 0.499, p = 0.4454
    0.04 mMBonferroni’s post hocCI(m): WT, 0.056 and 0.365; SD/SD, 0.003 and 0.312, p = 0.3294
    Fig. 2C
    Total sucrose intake
    WT and SA/SA miceNormal distributionPaired factorial ANOVAWT: n = 5, SA/SA: n = 5Genotype: F = 0.3578, p = 0.5663
    0.033% (w/v)Bonferroni’s post hocCI(m): WT, 0.442 and 1.558; SA/SA, 0.578 and 1.694, p = 0.4868
    0.1% (w/v)Bonferroni’s post hocCI(m): WT, 2.105 and 3.221; SA/SA, 2.258 and 3.375, p = 0.4345
    WT and SD/SD miceNormal distributionPaired factorial ANOVAWT: n = 5, SD/SD: n = 5Genotype: F = 0.2553, p = 0.6270
    0.033% (w/v)Bonferroni’s post hocCI(m): WT, 0.442 and 1.558; SD/SD, 0.214 and 1.329, p = 0.2855
    0.1% (w/v)Bonferroni’s post hocCI(m): WT, 2.359 and 3.475; SD/SD, 2.343 and 3.458, p = 0.9385
    Fig. 3
    Blood ethanol concentration
    WT and SA/SA miceNormal distributionPaired factorial ANOVAWT: n = 6, SA/SA: n = 5Genotype: F = 0.7817, p = 0.3996
    1 hTukey–Kramer post hocCI(m): WT, 2085.307 and 2908.035; SA/SA, 2108.846 and 3010.099, p = 0.6879
    3 hTukey–Kramer post hocCI(m): WT, 1063.610 and 1886.337; SA/SA, 1298.783 and 2200.036, p = 0.0947
    WT and SD/SD miceNormal distributionPaired factorial ANOVAWT: n = 5, SD/SD: n = 4Genotype: F = 0.1381, p = 0.7212
    1 hTukey–Kramer post hocCI(m): WT, 2115.538 and 3687.719; SD/SD, 2109.947 and 3867.699, p = 0.7418
    3 hTukey–Kramer post hocCI(m): WT, 1211.065 and 2783.246; SD/SD, 769.016 and 2526.767, p = 0.2089
    Fig. 4
    LORR
    WT and SA/SA miceNormal distributionWelch t test unpairedWT: n = 13, SA/SA: n = 13CI(d): –12.835 and 11.450, t = –0.118, p = 0.9072
    WT and SD/SD miceNormal distributionWelch t test unpairedWT: n = 16, SD/SD: n = 13CI(d): –6.790 and 28.367, t = 1.261, p = 0.2184
    Fig. 5A
    ROCK activity
    WT and SA/SA miceAssuming normalityWelch t test unpairedWT: n = 4, SA/SA: n = 4CI(d): 18.691 and 158.488, t = 3.305, p = 0.0228
    WT and SD/SD miceAssuming normalityWelch t test unpairedWT: n = 4, SD/SD: n = 4CI(d): –157.109 and 115.935, t = –0.421, p = 0.6956
    Fig. 5B
    ROCK protein level
    WT and SA/SA miceAssuming normalityWelch t test unpairedWT: n = 4, SA/SA: n = 4CI(d): –0.015 and 0.014, t = –0.141, p = 0.8929
    WT and SD/SD miceAssuming normalityWelch t test unpairedWT: n = 4, SD/SD: n = 4CI(d): –0.032 and 0.085, t = 1.248, p = 0.2782
    Fig. 5C
    ROCK activity/protein.
    WT and SA/SA miceAssuming normalityWelch t test unpairedWT: n = 4, SA/SA: n = 4CI(d): 194.255 and 925.975, t = 3.747, p = 0.0096
    WT and SD/SD miceAssuming normalityWelch t test unpairedWT: n = 4, SD/SD: n = 4CI(d): –1286.927 and 604.178, t = –0.937, p = 0.3929
    Fig. 5D
    Phosphorylated Akt
    WT and SA/SA miceNormal distributionWelch t test unpairedWT: n = 5, SA/SA: n = 5CI(d): –0.077 and –0.007, t = –2.751, p = 0.0256
    Table 2
    Body weight
    Fig. 1 (ethanol two-bottle choice test)
    WT and SA/SA mice (5%)Normal distributionWelch t test unpairedWT: n = 6, SA/SA: n = 7CI(d): –0.003 and 0.002, t = –0.512, p = 0.6229
    WT and SA/SA mice (10%)Normal distributionWelch t test unpairedWT: n = 5, SA/SA: n = 5CI(d): –0.002 and 0.009, t = 1.561, p = 0.1651
    WT and SD/SD mice (5%)Normal distributionWelch t test unpairedWT: n = 7, SD/SD: n = 6CI(d): –0.001 and 0.007, t = 1.907, p = 0.0874
    WT and SD/SD mice (10%)Normal distributionWelch t test unpairedWT: n = 8, SD/SD: n = 8CI(d): –0.003 and 0.003, t = 0.029, p = 0.9770
    Fig. 2 (taste preference test)
    WT/WT and SA/SA miceNormal distributionWelch t test unpairedWT: n = 5, SA/SA: n = 5CI(d): –0.005 and 0.007, t = 0.435, p = 0.6769
    WT/WT and SD/SD miceNormal distributionWelch t test unpairedWT: n = 5, SA/SA: n = 5CI(d): –0.004 and 0.009, t = 0.847, p = 0.4226
    Table 3
    Putative average ethanol consumption
    WT and SA/SA mice
    5% ethanolNormal distributionWelch t test unpairedWT: n = 6, SA/SA: n = 7CI(d): –2.223 and –0.209, t = –2.809, p = 0.0240
    10% ethanolNormal distributionWelch t test unpairedWT: n = 5, SA/SA: n = 5CI(d): –1.707 and –0.305, t = –3.309, p = 0.0107
    WT and SD/SD mice
    5% ethanolNormal distributionWelch t test unpairedWT: n = 7, SD/SD: n = 6CI(d): –1.692 and 0.833, t = –0.820, p = 0.4414
    10% ethanolNon-normal distributionMann–Whitney U testWT: n = 8, SD/SD: n = 8U = 26, p = 0.5286
    Table 4
    Putative average total liquid intake
    WT and SA/SA mice
    5% ethanolNormal distributionWelch t test unpairedWT: n = 6, SA/SA: n = 7CI(d): –19.2958 and 42.1470, t = 0.8822, p = 0.4074
    10% ethanolNormal distributionWelch t test unpairedWT: n = 5, SA/SA: n = 5CI(d): –12.7726 and 19.1717, t = 0.4800, p = 0.6468
    WT and SD/SD mice
    5% ethanolNormal distributionWelch t test unpairedWT: n = 7, SD/SD: n = 6CI(d): –29.0687 and 14.5159, t = –0.7377, p = 0.4766
    10% ethanolNormal distributionWelch t test unpairedWT: n = 8, SD/SD: n = 8CI(d): –25.3723 and 22.8610, t = –0.1142, p = 0.9111
    • CI(d), 95% confidence interval for the difference in population means, lower, and upper limits; CI(m), 95% confidence interval for the population means, lower, and upper limits.

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    Table 2.

    Summary of mean body weights during two-bottle choice tests for ethanol and for sucrose or quinine

    ExperimentMean body weight (kg)p value
    Fig. 1
    SA
    5% ethanol
    WT/WT0.028 ± 0.002 (n = 6)
    SA/SA0.028 ± 0.001 (n = 7)0.6229
    10% ethanol
    WT/WT0.031 ± 0.004 (n = 5)
    SA/SA0.028 ± 0.003 (n = 5)0.1651
    SD
    5% ethanol
    WT/WT0.030 ± 0.004 (n = 7)
    SD/SD0.027 ± 0.002 (n = 6)0.0874
    10% ethanol
    WT/WT0.028 ± 0.001 (n = 8)
    SD/SD0.028 ± 0.003 (n = 8)0.977
    Fig. 2
    Sucrose/quinine
    SA
    WT/WT0.036 ± 0.005 (n = 5)
    SA/SA0.035 ± 0.003 (n = 5)0.6769
    SD
    WT/WT0.033 ± 0.004 (n = 5)
    SD/SD0.031 ± 0.005 (n = 5)0.4226
    • Data are mean ± SD; p values are comparisons with WT/WT by unpaired two-tailed Welch’s t tests.

    • View popup
    Table 3.

    Daily average ethanol consumption in the two-bottle ethanol choice test

    Two-bottle ethanol choice testAverage ethanol consumption (g/kg/d)p value
    SA
    5% ethanol
    WT/WT0.952 ± 0.371 (n = 6)
    SA/SA2.168 ± 1.073 (n = 7)0.0240
    10% ethanol
    WT/WT1.193 ± 0.483 (n = 5)
    SA/SA2.199 ± 0.478 (n = 5)0.0107
    SD
    5% ethanol
    WT/WT0.945 ± 0.487 (n = 7)
    SD/SD1.375 ± 1.201 (n = 6)0.4414
    10% ethanol
    WT/WT1.294 ± 0.565 (n = 8)
    SD/SD1.352 ± 1.197 (n = 8)0.5286
    • Volitional ethanol intake was measured in a two-bottle choice test between tap water and 5% or 10% ethanol for three weeks in SA and SD mice. The measured values were divided by the number of days for each or the six measurements to obtain putative daily averages. Data are mean ± SD; p values are comparisons with WT/WT obtained by unpaired two-tailed Welch’s t tests, except for consumption of 10% ethanol in SD/SD mice, which was analyzed by Mann–Whitney U test.

    • View popup
    Table 4.

    Daily average total liquid intake in the two-bottle ethanol choice test

    Two-bottle ethanol choice testAverage total intake (g/kg/d)p value
    SA
    5% ethanol
    WT/WT139.3 ± 29.1 (n = 6)
    SA/SA127.9 ± 13.8 (n = 7)0.4074
    10% ethanol
    WT/WT132.1 ± 12.8 (n = 5)
    SA/SA128.9 ± 7.7 (n = 5)0.6468
    SD
    5% ethanol
    WT/WT109.2 ± 17.7 (n = 7)
    SD/SD116.4 ± 17.8 (n = 6)0.4766
    10% ethanol
    WT/WT118.8 ± 15.8 (n = 8)
    SD/SD120.0 ± 26.8 (n = 8)0.9111
    • Total liquid intake of a 5% or a 10% ethanol solution and water over a period of three weeks by SA and SD mice were measured in the two-bottle choice test. The measured values were divided by the number of days to obtain putative daily averages. Data are mean ± SD; p values are comparisons with WT/WT obtained using unpaired two-tailed Welch’s t tests.

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Nonphosphorylatable Src Ser75 Mutation Increases Ethanol Preference and Consumption in Mice
Goro Kato
eNeuro 28 March 2019, 6 (2) ENEURO.0418-18.2019; DOI: 10.1523/ENEURO.0418-18.2019

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Nonphosphorylatable Src Ser75 Mutation Increases Ethanol Preference and Consumption in Mice
Goro Kato
eNeuro 28 March 2019, 6 (2) ENEURO.0418-18.2019; DOI: 10.1523/ENEURO.0418-18.2019
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