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. Author manuscript; available in PMC: 2019 Jun 28.
Published in final edited form as: Cell. 2018 Jun 28;174(1):32–43.e15. doi: 10.1016/j.cell.2018.06.012

Figure 1. NMDA receptors in striatum are critical for learning heterogeneous action sequences.

Figure 1

(A) Operant chamber schematic. (B) Serial (Top) vs. hierarchical (Bottom) organization of LLRR sequence. (C and D) Example of typical wildtype mouse behavior on day 1 (C) and day 28 (D) of training. Top Panels: Left and right lever presses indicated by blue and red dashes, respectively, and aligned to magazine entry at time zero. Bottom Panels: Averaged left and right lever press rate indicated by blue and red lines, respectively. Insets show two representative sequences. (E) Development of a stereotypical action sequence across 28 days of training in a wildtype mouse. (F) Behavioral efficiency for control (n = 22 mice; main effect of training F4,84 = 81.71, P < 0.0001) and striatal NR1-KO mice (n = 5 mice; no effect of training F4,16 = 0.1531, P = 0.9588) across training (main effect of genotype F1,25 = 257, P < 0.0001). (G) Percentage of sequences beginning with the ‘LL’ subsequence (LL − −, ‘−’ denotes either ‘L’ or ‘R’ press in the sequence) or ending with the ‘RR’ subsequence (− − RR) for control (LL − − : main effect of training F4,84 = 18.27, P < 0.0001; − − RR: main effect of training F4,84 = 30.26, P < 0.0001) and striatal NR1-KO mice (LL − − : no effect of training F4,16 = 1.704, P = 0.1982; − − RR: no effect of training F4,16 = 0.6898, P = 0.6096) across training (LL − − : main effect of genotype F1,25 = 11.32, P = 0.0025; − − RR: main effect of genotype F1,25 = 26.93, P < 0.0001). (H) Percentage of sequences containing each appropriate element position for control (L − − − : main effect of training F4,84 = 24.17, P < 0.0001; − L − − : main effect of training F4,84 = 23.88, P < 0.0001; − − R − : main effect of training F4,84 = 29.97, P < 0.0001; − − − R: main effect of training F4,84 = 62.05, P < 0.0001; L − − − vs. − L − − : main effect of element position F1,21 = 9.616, P = 0.0054; first day of significant difference, Day 7, P = 0.0027; − − R − vs. − − − R: main effect of element position F1,21 = 41.56, P < 0.0001; first day of significant difference, Day 1, P = 0.013) and striatal NR1-KO mice (L − − − : no effect of training F4,16 = 1.747, P = 0.189; − L − − : no effect of training F4,16 = 1.077, P = 0.4006; − − R − : no effect of training F4,16 = 0.7249, P = 0.5877; − − − R: no effect of training F4,16 = 1.412, P = 0.275) across training (L − − − : main effect of genotype F1,25 = 22.77, P < 0.0001; − L − − : main effect of genotype F1,25 = 32.21, P < 0.0001; − − R − : main effect of genotype F1,25 = 27.01, P < 0.0001; − − − R: main effect of genotype F1,25 = 60.26, P < 0.0001). Data were analyzed using repeated-measures one-way or two-way ANOVA followed by Tukey or Sidak post-hoc comparisons, respectively. Error bars denote S.E.M., same for below unless stated otherwise. See also Figures S1 and S2.