Article Figures & Data
Figures
- Figure 1.
Skilled reaching performance improves with training. A, A single skilled reaching trial. (1) Rat breaks IR beam at the back of the chamber to request a sugar pellet (“beam break”). (2) Real-time analysis detects the hand breaching the reaching slot to trigger 300-fps video from 1 s before to 3.33 s after the trigger event (“video trigger”); (3) 2 s after the trigger event, the pellet delivery rod resets and the rat can initiate a new trial (“intertrial interval”). B, Average number of trials per day. Learners (green) and non-learners (pink) did not differ in the number of trials performed per day (linear mixed model: effect of group: t(28) = 0.49, p = 0.63). However, both groups increased the number of trials performed over days (linear mixed model: effect of day: t(124) = 3.58, p = 4.88 × 10−4; group × day interaction: t(124) = −0.47, p = 0.64). Black line represents the average number of trials per day for both groups combined. C, Average first attempt success rate per day. Although both groups had similar success rates on day 1, only learners increased their success rates over days (linear mixed model: effect of group: t(25) = 0.44, p = 0.66; effect of day: t(123) = 5.55, p = 1.70 × 10−7; group × day interaction: t(123) = −4.78, p = 4.98 × 10−6. Error bars in B, C represent SEM. Extended Data Figure 1-1 shows individual rat data for each group and additional performance measures (number of attempts per trial and breakdown of reach outcomes across days); ***p < 0.001 for the day term in the linear mixed model in B and ###p < 0.001 for the group × day interaction in the linear mixed model in C. Data and code to generate this figure are contained in Extended Data 1, 2.
- Figure 2.
Refinement of “gross” forelimb movements. A, All 3D reaching trajectories on days 1 and 10 from an exemplar rat. Colored lines represent individual trials and black lines represent average trajectories of the hand and digit tips. Sugar pellet (black dot) is at (0,0,0). B, Hand trajectories are separated into “reach” and “grasp” components. The reach begins when the digits are first visible outside the reaching box and ends when the second digit begins to retract. The grasp begins when the second digit begins to flex and ends when the second digit is maximally flexed after grasp start. C, Average hand trajectory variability for the reach and grasp components represented as the mean distance from the average trajectory (mm). Trajectory variability did not differ significantly between learners and non-learners for either component (linear mixed model: effect of group: reach: t(23) = 0.53, p = 0.60; grasp: t(26) = −0.08, p = 0.94; group × day interaction: reach: t(124) = 0.37, p = 0.71; grasp: t(124) = 1.08, p = 0.28). Trajectory variability of the grasp, but not reach, decreased significantly over days for both groups (linear mixed model: effect of day: reach: t(124) = −1.85, p = 0.07; grasp: t(124) = −2.75, p = 6.91 × 10−3). Individual rat data are shown in Extended Data Figure 2-1. D, Moving average of hand trajectory variability for the reach component (black) and first attempt success rate (green learners and pink non-learners) within individual days. Days 2, 4, 5, 6, and 8 had significant negative correlations between trajectory variability and success rate for the learners group (day 2: r = −0.55, p = 1.70 × 10−3; day 4: r = −0.68, p = 4.25 × 10−5; day 5: r = −0.53, p = 2.50 × 10−3; day 6: r = −0.82, p = 2.16 × 10−8; day 8: r = −0.70, p = 1.51 × 10−5). Days 1, 4, 6, 8, and 10 had significant negative correlations between trajectory variability and success rate for the non-learners group (day 1: r = −0.88, p = 1.78 × 10−10; day 4: r = −0.67, p = 4.33 × 10−5; day 6: r = −0.38, p = 0.04; day 8: r = −0.42, p = 0.02; day 10: r = −0.83, p = 1.53 × 10−8). E, Average reach duration for learners (green) and non-learners (pink). Linear mixed model: effect of group: t(21) = 0.29, p = 0.78; effect of day: t(124) = −1.03, p = 0.31; group × day interaction: t(124) = −0.11, p = 0.91. Individual rat data are shown in Extended Data Figure 2-1. F, Average maximum reach velocity (mm/s) for learners (green) and non-learners (pink). Linear mixed model: effect of group: t(15) = 0.16, p = 0.87; effect of day: t(124) = −0.07, p = 0.94; group × day interaction: t(124) = 1.02, p = 0.31. Individual rat data are shown in Extended Data Figure 2-1. Error bars in C, E, F, and shaded colored areas in D represent SEM; *p < 0.05, **p < 0.01, ***p < 0.001 for a negative correlation between success rate and trajectory variability in D; **p < 0.05 for the day term in the linear mixed model in C. Data and code to generate this figure are contained in Extended Data 1, 2.
- Figure 3.
Reach endpoint accuracy improves with training. A, X, Y, and Z reach endpoints are measured in reference to the pellet position. Top, Mirror view of the dorsal surface of the hand. Bottom, Direct view from the camera. The inset shows the mirror view of the palmar surface of the hand to demonstrate how reach endpoint of the second digit tip in the Z direction (zdigit2) is measured. The end of the reach is defined as the moment zdigit2 begins to decrease (the digit tip moves back toward the box). B, Average reach endpoints of the hand (left) and digit 2 (right) in the X, Y, and Z directions. Pellet is at (0,0,0). Reach endpoint in the X direction increased over days for learners but not non-learners. Linear mixed model (hand): effect of group: t(18) = 1.30, p = 0.21; effect of day: t(124) = 2.13, p = 0.04; group × day interaction: t(124) = −2.40, p = 0.02; digit 2: effect of group: t(17) = 1.80, p = 0.09; effect of day: t(124) = 3.12, p= 2.25 × 10−3; group × day interaction: t(124) = −3.16, p = 1.97 × 10−3. Reach endpoint of the hand in the Y plane decreased with training for both groups. Linear mixed model (hand): effect of group: t(18) = −0.94, p = 0.36; effect of day: t(124) = −2.45, p = 0.02; group × day interaction: t(124) = 1.45, p = 0.15; digit 2: effect of group: t(19) = −0.90, p = 0.38; effect of day: t(124) = −1.80, p = 0.07; group × day interaction: t(124) = 1.16, p = 0.25. Reach endpoint in the Z plane did not change significantly with training for either group. Linear mixed model (hand): effect of group: t(23) = −0.38, p = 0.71; effect of day: t(124) = 1.73, p = 0.09; group × day interaction: t(124) = −1.23, p = 0.22; digit 2: effect of group: t(19) = −0.63, p = 0.53; effect of day: t(124) = 0.99, p = 0.33; group × day interaction: t(124) = −0.58; p = 0.57. Extended Data Figure 3-1 shows individual rat data. C, 3D covariance of hand and digit reach endpoints across all days from an exemplar rat. Sugar pellet (*) is at (0,0,0). D, Average determinant of the covariance matrix (generalized variance) of reach endpoints for the hand and digits. Top row shows endpoint variability of “raw” digit and hand position data (linear mixed model: effect of day: digit 1: t(124) = −1.97, p = 0.05; digit 2: t(124) = −2.25, p = 0.03; digit 3: t(124) = −1.85, p = 0.07; digit 4: t(124) = −1.78, p = 0.08; hand: t(124) = −2.27, p = 0.02). Bottom row shows endpoint variability of digit positions subtracted from hand position (linear mixed model: effect of day: digit 1: t(124) = −0.87, p = 0.39; digit 2: t(124) = −2.23, p = 0.03; digit 3: t(136) = −1.64, p = 0.10; digit 4: t(124) = −0.67, p = 0.51). Extended Data Figure 3-2 shows individual rat data. E, Average percentage of frames from reach start to grasp end that were mislabeled by DeepLabCut for the hand (dark blue) and digits 1–4 (light blue, pink, yellow, and green, respectively) on days 1 and 10. The percentage of mislabeled frames did not differ between parts (linear mixed model: effect of part: t(123) = −0.05, p = 0.96) or days overall (linear mixed model: effect of day: t(123) = 0.98, p = 0.33). The percentage of mislabeled frames increased significantly between day 1 and day 10 for digit 4 only (linear mixed model: part × day interaction: hand: t(117) = −0.79, p = 0.43; digit 1: t(117) = 1.07, p = 0.29; digit 2: t(117) = 0.91, p = 0.37; digit 3: t(117) = 1.82, p = 0.07; digit 4: t(117) = 2.71, p = 7.74 × 10−3). Error bars in B, D, E, represent SEM; *p < 0.05 for the day term, #p < 0.05, ##p < 0.01 for the group × day interaction in the linear mixed model in B, D; #p < 0.05 for the part × day interaction in E. Data and code to generate this figure are contained in Extended Data 1, 2.
- Figure 4.
Refinement of “fine” digit movements. A, Average digit 2 trajectory variability for the reach and grasp components represented as the mean distance from the average trajectory (mm). Trajectory variability significantly decreased over days during the grasp, but not reach, component (linear mixed model: effect of day: reach: t(124) = −1.70, p = 0.09; grasp: t(124) = −3.92, p = 1.46 × 10−4). Trajectory variability of the grasp was significantly lower for learners than non-learners in the later training days (group × day interaction: reach: t(124) = −0.01, p = 0.99; grasp: t(124) = 2.29, p = 0.02). Individual rat data are shown in Extended Data Figure 4-1. B, Moving average of digit 2 trajectory variability for the reach component for learners (green) and non-learners (pink) within individual days. C, Grasp aperture (a) is the Euclidian distance between the first and fourth digit tips. D, Average grasp aperture at reach end for learners (green) and non-learners (pink). Linear mixed model: effect of group: t(15) = −0.81, p = 0.43; effect of day: t(124) = 0.97, p = 0.34; group × day interaction: t(124) = 0.36, p = 0.72. Individual rat data are shown in Extended Data Figure 4-1. E, Moving average of grasp aperture at reach end within days for learners (green) and non-learners (pink). F, Average aperture variance at reach end. Linear mixed model: effect of group: t(22) = 0.96, p = 0.35; effect of day: t(124) = −1.53, p = 0.13; group × day interaction: t(124) = 0.90, p = 0.37. Individual rat data are shown in Extended Data Figure 4-1. G, Hand orientation is the angle (θ) between a line connecting the first and fourth digit tips and the floor. H, Average hand orientation at reach end increased over days but did not differ between groups. Linear mixed model: effect of group: t(15) = 0.78, p = 0.45; effect of day: t(124) = 2.72, p = 7.42 × 10−3; group × day interaction: t(124) = −1.54, p = 0.13. Individual rat data are shown in Extended Data Figure 4-1. I, Moving average of hand orientation at reach end within individual days for learners (green) and non-learners (pink). J, Average hand orientation MRL at reach end. Linear mixed model: effect of group: t(61) = 0.80, p = 0.43; effect of day: t(124) = 1.97, p = 0.05; group × day interaction: t(124) = −0.98, p = 0.33. Individual rat data are shown in Extended Data Figure 4-1. K, Digit flexion is the angle (δ) between a line joining the second MCP joint and hand dorsum; and a line joining the second MCP joint and the tip of digit 2. L, Average digit flexion at reach end. Linear mixed model: effect of group: t(16) = −0.68, p = 0.51; effect of day: t(124) = −1.49, p = 0.14; group × day interaction: t(124) = 0.98, p = 0.33. Individual rat data are shown in Extended Data Figure 4-1. M, Moving average of digit flexion at reach end within individual days for learners (green) and non-learners (pink). N, Average digit flexion MRL at reach end. Linear mixed model: effect of group: t(26) = −1.00, p = 0.33; effect of day: t(124) = 1.02, p = 0.31; group × day interaction: t(124) = −0.01, p = 0.99. Individual rat data are shown in Extended Data Figure 4-1. Error bars in A, D, F, H, J, L, N and shaded areas in B, E, I, M represent SEM; **p < 0.01 and ***p < 0.001 for the day term in the linear mixed model in A, H; #p < 0.05 for the group × day interaction in A. Data and code to generate this figure are contained in Extended Data 1, 2.
- Figure 5.
Coordination between digit movements and hand advancement improves with training. A, Mean aperture as a function of hand advancement (zdigit2, pellet at zdigit2 = 0) across 10 training days for learners (green) and non-learners (pink). D1–2, D3–4, … represent days 1–2, days 3–4, etc. Dashed line indicates the zdigit2 coordinate (+3 mm) where data are sampled in B. B, Average grasp aperture at the zdigit2 coordinate (+3 mm) indicated by the dashed line in A as a function of day number. Linear mixed model: effect of group: t(30) = −0.53, p = 0.60; effect of day: t(111) = −0.17, p = 0.86; group × day interaction: t(111) = 1.35, p = 0.18. Extended Data Figure 5-1 shows individual rat data. C, Mean hand orientation (θ) as a function of hand advancement across all days for learners (green) and non-learners (pink). Dashed line indicates the zdigit2 coordinate (+3 mm) where data are sampled in D. D, Average hand orientation (θ) at the zdigit2 coordinate (+3 mm) indicated by the dashed lines in C across days. Linear mixed model: effect of group: t(18) = 1.11, p = 0.28; effect of day: t(111) = 2.20, p = 0.03; group × day interaction: t(111) = −2.08, p = 0.04. Extended Data Figure 5-1 shows individual rat data. E, Mean digit flexion (δ) as a function of hand advancement for learners (green) and non-learners (pink). Dashed line indicates the zdigit2 coordinate (+3 mm) where data are sampled in F. F, Average digit flexion (δ) at the zdigit2 coordinate (+3 mm) indicated by the dashed lines in E across days. Linear mixed model: effect of group: t(19) = −0.18, p = 0.86; effect of day: t(111) = −0.05, p = 0.96; group × day interaction: t(111) = −0.52, p = 0.60. Extended Data Figure 5-1 shows individual rat data. Error bars in B, D, F represent SEM; *p < 0.05 for the day term and #p < 0.05 for the group × day interaction in the linear mixed model in D. Data and code to generate this figure are contained in Extended Data 1, 2.
- Figure 6.
Kinematic measures separated by reach success or failure. A, Average hand trajectory variability (mean distance from the average trajectory, mm) for the reach component in learners (green) and non-learners (pink). Trajectory variability was significantly higher for failed reaches (“miss,” black) than successful reaches (“hit,” green/pink) for non-learners, but not for learners. Linear mixed model (learners): effect of outcome: t(72) = 1.26, p = 0.21; outcome × day interaction: t(72) = −0.46, p = 0.64. Linear mixed model (non-learners): effect of outcome: t(183) = 4.44, p = 1.56 × 10−5; outcome × day interaction: t(183) = −1.25, p = 0.21. Individual rat data are shown in Extended Data Figure 6-1. B, Average hand trajectory variability (mean distance from the average trajectory, mm) for the grasp component in learners (green) and non-learners (pink). Trajectory variability was significantly higher for failed reaches (“miss,” black) than successful reaches (“hit,” green/pink) for both groups. Linear mixed model (learners): effect of outcome: t(72) = 4.77, p = 9.34 × 10−6; outcome × day interaction: t(72) = −2.23, p = 0.03. Linear mixed model (non-learners): effect of outcome: t(183) = 5.58, p = 8.47 × 10−8; outcome × day interaction: t(183) = −1.02, p = 0.31. Individual rat data are shown in Extended Data Figure 6-1. C, Average reach endpoint of digit 2 in the X, Y, and Z directions for learners (top) and non-learners (bottom). Reach endpoint in the X direction was significantly different for successful (green/pink) versus failed (black) reaches for learners but not non-learners. Linear mixed model (learners): effect of outcome: t(72) = −2.31, p = 0.02; outcome × day interaction: t(72) = 1.94, p = 0.06. Linear mixed model (non-learners): effect of outcome: t(185) = −1.53, p = 0.13; outcome × day interaction: t(185) = −0.03, p = 0.97. Reach endpoint in the Y direction did not differ between successful and failed reaches for either group. Linear mixed model (learners): effect of outcome: t(72) = −0.95, p = 0.35; outcome × day interaction: t(72) = 0.23, p = 0.82. Linear mixed model (non-learners): effect of outcome: t(185) = −1.49, p = 0.14; outcome × day interaction: t(185) = 0.77, p = 0.44. Failed reaches were significantly shorter (Z direction) than successful reaches for both groups, although the Z endpoint did not differ between successful and failed reaches in later training days for learners. Linear mixed model (learners): effect of outcome: t(72) = 3.52, p = 7.48 × 10−4; outcome × day interaction: t(72) = −1.86, p = 0.07. Linear mixed model (non-learners): effect of outcome: t(185) = 4.94, p = 1.76 × 10−6; outcome × day interaction: t(185) = −1.53, p = 0.13. Individual rat data are shown in Extended Data Figure 6-2. D, Average determinant of the covariance matrix (generalized variance) of reach endpoints for the hand across days for learners (green) and non-learners (pink). There was significantly greater variability in reach endpoint for failed (black) reaches than successful (green/pink) reaches for both groups, although endpoint variability did not differ between successful and failed reaches in later training days for learners. Linear mixed model (learners): effect of outcome: t(72) = 2.67, p = 9.26 × 10−3; outcome × day interaction: t(72) = −1.95, p = 0.05; non-learners: effect of outcome: t(185) = 2.63, p = 9.34 × 10−3; outcome × day interaction: t(185) = −0.67, p = 0.50. Individual rat data are shown in Extended Data Figure 6-2. E, Average determinant of the covariance matrix (generalized variance) of reach endpoints for digit 2 across days for learners (green) and non-learners (pink). There was significantly greater variability in reach endpoint for failed (black) reaches than successful (green/pink) reaches for both groups, although endpoint variability did not differ between successful and failed reaches in later training days for learners. Linear mixed model (learners): effect of outcome: t(72) = 2.64, p = 0.01; outcome × day interaction: t(72) = −1.92, p = 0.06; non-learners: effect of outcome: t(185) = 2.60, p = 0.01; outcome × day interaction: t(185) = −0.25, p = 0.80. Individual rat data are shown in Extended Data Figure 6-2. Error bars in A–E represent SEM; *p < 0.05 for the outcome term in the linear mixed model in C, E, **p < 0.01 for the outcome term in the linear mixed model in D, ***p < 0.001 for the outcome term in the linear mixed model in A–C; #p < 0.05 for the outcome × day interaction in the linear mixed model in B. Summaries of key metrics for every trial performed by each rat, separated by trial outcomes, are shown in Extended Data Figures 6-3, 6-4, 6-5, 6-6, 6-7, 6-8, 6-9, 6-10, 6-11, 6-12, 6-13, 6-14, 6-15, 6-16. Data and code to generate this figure are contained in Extended Data 1, 2.
Tables
Figure Data structure Type of test Sample
sizeStatistical data Fig. 1B Effect of group on number of trials
Effect of day on number of trials
Group × day interactionLinear mixed model n = 14 t = 0.49, df = 28.48, p = 0.63
t = 124, df = 3.58, p = 4.88 × 10−4
t = 124, df = −0.47, p = 0.64Fig. 1C Effect of group on first reach success rate
Effect of day on first reach success rate
Group × day interactionLinear mixed model n = 14 t = 0.442, df = 25, p = 0.66
t = 5.55, df = 123, p = 1.70 × 10−7
t = −4.78, df = 123, p = 4.98 × 10−6Extended Data Fig. 1-1C Effect of group on number of reach attempts
Effect of day on number of reach attempts
Group × day interactionLinear mixed model n = 14 t = 0.81, df = 19, p = 0.43
t = −4.12, df = 124, p = 6.88 × 10−5
t = 1.37, df = 124, p = 0.17Extended Data Fig. 1-1D Effect of day on performance outcomes, learners
Effect of day on performance outcomes, non-learnersLinear mixed model n = 4 No pellet: t = −0.11, df = 35, p = 0.91
First success: t = 4.96, df = 35, p = 1.83 × 10−5
Multiple success: t = −2.53, df = 35, p = 0.02
Drop in box: t = 0.42, df = 35, p = 0.68
Pellet knock off: t = −4.59, df = 35, p =5.46 × 10−5
Tongue: t = 0, df = 35, p = 1
Trigger error: t = −0.52, df = 38, p = 0.61
Pellet remained: t = −0.78, df = 38, p = 0.44
Non-preferred hand: t = −0.87, df = 38, p = 0.39
Tongue and hand: t = 0, df = 38, p = 1
Hand through slot: t = −0.78, df = 38, p = 0.39
No pellet: t = −0.62, df = 98, p = 0.54
First success: t = 0.51, df = 88, p = 0.61
Multiple success: t = −0.79, df = 89, p = 0.43
Drop in box: t = 0.36, df = 89, p = 0.72
Pellet knock off: t = 0.49, df = 89, p = 0.62
Tongue: t = 0, df = 98, p = 1
Trigger error: t = −1.37, df = 98, p = 0.18
Pellet remained: t = −0.95, df = 98, p = 0.35
Non-preferred hand: t = −0.87, df = 98, p = 0.39
Tongue and hand: t = 0, df = 98, p = 1
Hand through slot: t = 0.01, df = 98, p = 0.99Fig. 2C Effect of group on paw trajectory variability
Effect of day on paw trajectory variability
Group × day interactionLinear mixed model n = 14 Reach: t = 0.53, df = 23, p = 0.60
Grasp: t = −0.08, df = 26, p = 0.94
Reach: t = −1.85, df = 124, p = 0.07
Grasp: t = −2.75, df = 124, p = 6.91 × 10−3
Reach: t = 0.37, df = 124, p = 0.71
Grasp: t = 1.08, df = 124, p = 0.28Fig. 2D Negative correlation between trajectory variability and success rate, learners
Negative correlation between trajectory variability and success rate, non-learnersLinear correlation n = 4
n = 10Session 1: r = 0.73, p = 5.33 × 10−6
Session 2: r = −0.55, p = 1.70 × 103
Session 3: r = −0.03, p = 0.86
Session 4: r = −0.68, p = 4.25 × 10−5
Session 5: r = −0.53, p = 2.50 × 10−3
Session 6: r = −0.82, p = 2.16 × 10−8
Session 7: r = 0.15, p = 0.44
Session 8: r = −0.70, p = 1.51 × 10−5
Session 9: r = 0.39, p = 0.03
Session 10: r = −0.24, p = 0.20
Session 1: r = −0.88, p = 1.78 × 10−10
Session 2: r = −0.02, p = 0.91
Session 3: r = 0.02, p = 0.91
Session 4: r = −0.67, p = 4.33 × 10−5
Session 5: r = −0.33, p = 0.08
Session 6: r = −0.38, p = 0.04
Session 7: r = 0.67, p = 4.33 × 10−5
Session 8: r = −0.42, p = 0.02
Session 9: r = −0.07, p = 0.73
Session 10: r = −0.83, p = 1.53 × 10−8Fig. 2E Effect of group on reach duration
Effect of day on reach duration
Group × day interactionLinear mixed model n = 14 t = 0.29, df = 21, p = 0.78
t = −1.03, df = 124, p = 0.31
t = −0.11, df = 124, p = 0.91Fig. 2F Effect of group on reach velocity
Effect of day on reach duration
Group × day interactionLinear mixed model n = 14 t = 0.16, df = 15, p = 0.87
t = −0.07, df = 124, p = 0.94
t = 1.02, df = 124, p = 0.31Fig. 3B Effect of group on reach endpoint
Effect of day on reach endpoint
Group × day interactionLinear mixed model n = 14 Paw x: t =1.30, df = 18, p = 0.21
Paw y: t = −0.94, df = 18, p = 0.36
Paw z: t = −0.38, df = 124, p = 0.71
Digit 2 ×: t = 1.80, df = 17, p = 0.09
Digit 2 year: t = −0.90, df = 19, p = 0.38
Digit 2 z: t = −0.63, df = 19, p = 0.53
Paw x: t = 2.13, df = 124, p = 0.04
Paw y: t = −2.45, df = 124, p = 0.02
Paw z: t = 1.73, df = 124, p = 0.09
Digit 2 ×: t = 3.12, df = 124, p = 2.25 × 103
Digit 2 year: t = −1.80, df = 124, p = 0.07
Digit 2 z: t = 0.99, df = 124, p = 0.33
Paw x: t = −2.40, df = 124, p = 0.02
Paw y: t = 1.45, df = 124, p = 0.15
Paw z: t = −1.23, df = 124, p = 0.22
Digit 2 ×: t = −3.16, df = 124, p = 1.97 × 103
Digit 2 year: t = 1.16, df = 124, p = 0.25
Digit 2 z: t = −0.58, df = 124, p = 0.57Fig. 3D Effect of group on endpoint variability, raw data
Effect of day on endpoint variability, raw data
Group × day interaction, raw data
Effect of group on endpoint variability, subtracted position
Effect of day on endpoint variability, subtracted position
Group × day interaction, subtracted positionLinear mixed model n = 14 Hand: t = −0.95, df = 51, p = 0.35
Digit 1: t = −0.71, df = 56, p = 0.48
Digit 2: t = −0.86, df = 45, p = 0.39
Digit 3: t = −0.42, df = 49, p = 0.67
Digit 4: t = −0.29, df = 48, p = 0.78
Hand: t = −2.27, df = 124, p = 0.02
Digit 1: t = −1.97, df = 124, p = 0.05
Digit 2: t = −2.25, df = 124, p = 0.03
Digit 3: t = −1.85, df = 124, p = 0.07
Digit 4: t = −1.78, df = 124, p = 0.08
Hand: t = 1.39, df = 124, p = 0.17
Digit 1: t = 1.15, df = 124, p = 0.25
Digit 2: t = 1.42, df = 124, p = 0.16
Digit 3: t = 0.90, df = 124, p = 0.37
Digit 4: t = 0.89, df = 124, p = 0.37
Digit 1: t = 0.13, df = 78, p = 0.89
Digit 2: t = −1.61, df = 112, p = 0.11
Digit 3: t = −0.75, df = 136, p = 0.45
Digit 4: t = 0.62, df = 107, p = 0.54
Digit 1: t = −0.87, df = 124, p = 0.39
Digit 2: t = −2.23, df = 124, p = 0.03
Digit 3: t = −1.64, df = 136, p = 0.10
Digit 4: t = −0.67, df = 124, p = 0.51
Digit 1: t = 0.64, df = 124, p = 0.52
Digit 2: t = 1.79, df = 124, p = 0.08
Digit 3: t = 1.21, df = 136, p = 0.23
Digit 4: t = 0.03, df = 124, p = 0.98Fig. 3E Effect of part on % mislabeled frames
Effect of day on % mislabeled frames
Part × day interactionLinear mixed model n = 14 t = −0.05, df = 123, p = 0.96
t = 0.98, df = 123, p = 0.33
Hand: t = −0.79, df = 117, p = 0.43
Digit 1: t = 1.07, df = 117, p = 0.29
Digit 2: t = 0.91, df = 117, p = 0.37
Digit 3: t = 1.82, df = 117, p = 0.07
Digit 4: t = 2.71, df = 117, p = 7.74 × 10−3Fig. 4A Effect of group on digit 2 trajectory variability
Effect of day on digit 2 trajectory variability
Group × day interactionLinear mixed model n = 14 Reach: t = 1.39, df = 23, p = 0.18
Grasp: t = −0.60, df = 27, p = 0.55
Reach: t = −1.69, df = 124, p = 0.09
Grasp: t = −3.92, df = 124, p = 1.46 × 10−4
Reach: t = −0.01, df = 124, p = 0.99
Grasp: t = 2.29, df = 124, p = 0.02Fig. 4D Effect of group on aperture
Effect of day on aperture
Group × day interactionLinear mixed model n = 14 t = −0.81, df = 15, p = 0.43
t = 0.97, df = 124, p = 0.34
t = 0.36, df = 124, p = 0.72Fig. 4F Effect of group on aperture variance
Effect of day on aperture variance
Group × day interactionLinear mixed model n = 14 t = 0.96, df = 22, p = 0.35
t = −1.53, df = 124, p = 0.13
t = 0.90, df = 124, p = 0.37Fig. 4H Effect of group on hand orientation
Effect of day on hand orientation
Group × day interactionLinear mixed model n = 14 t = 0.78, df = 15, p = 0.45
t = 2.72, df = 124, p = 7.42 × 10−3
t = −1.54, df = 124, p = 0.13Fig. 4J Effect of group on orientation variance
Effect of day on orientation variance
Group × day interactionLinear mixed model n = 14 t = 0.80, df = 61, p = 0.43
t = 1.97, df = 124, p = 0.05
t = −0.98, df = 124, p = 0.33Fig. 4L Effect of group on digit flexion
Effect of day on digit flexion
Group × day interactionLinear mixed model n = 14 t −0.68, df = 16, p = 0.51
t = −1.49, df = 124, p = 0.14
t = 0.98, df = 124, p = 0.33Fig. 4N Effect of group on flexion variance
Effect of day on flexion variance
Group × day interactionLinear mixed model n = 14 t = −1.00, df = 26, p = 0.33
t = 1.02, df = 124, p = 0.31
t = −0.01, df = 124, p = 0.99Fig. 5B Effect of group on aperture
Effect of day on aperture
Group × day interactionLinear mixed model n = 14 t = −0.53, df = 30, p = 0.60
t = −0.17, df = 111, p = 0.86
t = 1.35, df = 111, p = 0.18Fig. 5D Effect of group on orientation
Effect of day on orientation
Group × day interactionLinear mixed model n = 14 t = 1.11, df = 18, p = 0.28
t = 2.20, df = 111, p = 0.03
t = −2.08, df = 111, p = 0.04Fig. 5F Effect of group on flexion
Effect of day on flexion
Group × day interactionLinear mixed model n = 14 t = −0.184, df = 19, p = 0.86
t = −0.05, df = 111, p = 0.96
t = −0.52, df = 111, p = 0.60Fig. 6A Effect of outcome on reach trajectory variability
Effect of day on reach trajectory variability
Outcome × day interactionLinear mixed model n = 4
n = 10
n = 4
n = 10
n = 4
n = 10Learners: t = 1.26, df = 72, p = 0.21
Non-learners: t = 4.44, df = 183, p = 1.56 × 10−5
Learners: t = −1.84, df = 72, p = 0.07
Non-learners: t = −2.01, df = 183, p = 0.046
Learners: t = −0.46, df = 72, p = 0.64
Non-learners: t = −1.25, df = 183, p = 0.21Fig. 6B Effect of outcome on grasp trajectory
variability
Effect of day on grasp trajectory variability
Outcome × day interactionLinear mixed model n = 4
n = 10
n = 4
n = 10
n = 4
n = 10Learners: t = 4.77, df = 72, p = 9.34 × 10−6
Non-learners: t = 5.58, df = 183, p = 8.47 × 10−8
Learners: t = −0.30, df = 72, p = 0.77
Non-learners: t = −2.88, df = 183, p = 4.42 × 103
Learners: t = −2.23, df = 72, p = 0.03
Non-learners: t = −1.02, df = 183, p = 0.31Fig. 6C Effect of outcome on reach endpoint
Effect of day on reach endpoint
Outcome × day interactionLinear mixed model n = 4
n = 10
n = 4
n = 10
n = 4Learners X: t = −2.31, df = 72, p = 0.02
Learners Y: t = −0.95, df = 72, p = 0.35
Learners Z: t = 3.52, df = 72, p =7.48 × 10−4
Non-learners X: t = −1.53, df = 185, p = 0.13
Non-learners Y: t = −1.49, df = 185, p = 0.14
Non-learners Z: t = 4.94, df = 185, p = 1.76 × 10−6
Learners X: t = 0.95, df = 72, p = 0.34
Learners Y: t = 1.54, df = 72, p = 0.13
Learners Z: t = 1.31, df = 72, p = 0.20
Non-learners X: t = −1.46, df = 185, p = 0.15
Non-learners Y: t = 0.59, df = 185, p = 0.56
Non-learners Z: t = 0.76, df = 185, p = 0.45
Learners X: t = 1.94, df = 72, p = 0.06
Learners Y: t = 0.23, df = 72, p = 0.82
Learners Z: t = −1.86, df = 72, p = 0.07
Non-learners X: t = −0.03, df = 185, p = 0.97
Non-learners Y: t = 0.77, df = 185, p = 0.44
Non-learners Z: t = −1.53, df = 185, p = 0.13Fig. 6D Effect of outcome on hand endpoint variability
Effect of day on hand endpoint variability
Outcome × day interactionLinear mixed model n = 4
n = 10
n = 4
n = 10
n = 4
n = 10Learners: t = 2.67, df = 72, p = 9.26 × 10−3
Non-learners: t = 2.63, df = 185, p = 9.34 × 10−3
Learners: t = −0.06, df = 72, p = 0.96
Non-learners: t = −0.26, df = 185, p = 0.80
Learners: t = −1.95, df = 72, p = 0.05
Non-learners: t = −0.67, df = 185, p = 0.50Fig. 6E Effect of outcome on digit 2 endpoint variability
Effect of day on digit 2 endpoint variability
Outcome × day interactionLinear mixed model n = 4
n = 10
n = 4
n = 10
n = 4
n = 10Learners: t = 2.64, df = 72, p = 0.01
Non-learners: t = 2.60, df = 185, p = 0.01
Learners: t = −0.14, df = 72, p = 0.89
Non-learners: t = −0.22, df = 185, p = 0.83
Learners: t = −1.92, df = 72, p = 0.06
Non-learners: t = −0.25, df = 185, p = 0.80Rat Sex h p χ2 stat df Positive
change?Learner/non-learner A M 0 0.89 0.02 1 Yes Non-learner B M 1 0.04 4.45 1 Yes Learner C M 0 0.28 1.15 1 Yes Non-learner D M 0 0.26 1.28 1 Yes Non-learner E F 0 0.21 1.57 1 Yes Non-learner F M 1 4.58e-11 43.35 1 Yes Learner G M 0 0.63 0.24 1 Yes Non-learner H F 0 0.22 1.54 1 Yes Non-learner I F 1 0.02 5.52 1 Yes Learner J F 1 4.98e-4 12.13 1 Yes Learner K F 0 0.46 0.54 1 No Non-learner L M 1 0.02 5.38 1 No Non-learner M M 0 0.87 0.03 1 Yes Non-learner N M 0 0.91 0.01 1 No Non-learner
Extended Data Figure 1-1
Additional task performance measures. A, Individual rat data for average number of trials performed per day for learners (left) and non-learners (right). Each colored line represents an individual rat. Colors represent the same rat across all individual rat data figures within groups. B, Individual rat data for first reach success rate for learners (left) and non-learners (right). C, Number of reach attempts per trial. Both learners (left) and non-learners (right) decreased the number of reach attempts performed per trial over days (linear mixed model: effect of group: t(19) = 0.81, p = 0.43; effect of day: t(124) = –4.12, p = 6.88 × 10−5; group × day interaction: t(124) = 1.37, p = 0.17. Black line represents averaged data. Colored lines represent individual rats. D, Breakdown of trial outcomes by day for learners (green) and non-learners (for definitions of outcomes, see Materials and Methods, Number of trials and success rate). The percentage of “first success” increased over days only for learners (linear mixed model: effect of day: learners: t(35) = 4.96, p = 1.83 × 10−5; non-learners: t(88) = 0.51, p = 0.61). “Multiple success” outcomes decreased over days for learners but not non-learners (linear mixed model: effect of day: learners: t(35) = –2.53, p = 0.02; non-learners: t(89) = –0.79, p = 0.43). Similarly, “pellet knocked off” outcomes decreased for learners but not non-learners over days (linear mixed model: effect of day: learners: t(35) = –4.59, p = 5.46 × 10−5; non-learners: t(89) = 0.49, p = 0.62). All other outcomes were consistent across days for both groups. Linear mixed model: effect of day (learners): no pellet: t(35) = –0.11, p = 0.91; drop in box; t(35) = 0.42, p = 0.68; tongue: t(35) = 0; p = 1; trigger error: t(38) = –0.52, p = 0.61; pellet remained: t(38) = –0.78, p = 0.44; non-preferred hand: t(38) = –0.87, p = 0.39; tongue and hand: t(38) = 0, p = 1; hand through slot: t(38) = –0.78, p = 0.44. Linear mixed model: effect of day (non-learners): no pellet: t(98) = –0.62, p = 0.54; drop in box; t(89) = 0.36, p = 0.72; tongue: t(98) = 0; p = 1; trigger error: t(98) = –1.37, p = 0.18; pellet remained: t(98) = –0.95, p = 0.35; non-preferred hand: t(98) = –0.87, p = 0.39; tongue and hand: t(98) = 0, p = 1; hand through slot: t(98) = 0.01, p = 0.99; ***p < 0.001 for the day term in the linear mixed model in C, D; *p < 0.05 for the day term in the linear mixed model in D. Data and code to generate this figure are contained in Extended Data 1, 2. Download Figure 1-1, TIF file.
Extended Data 1
MATLAB and Python code for kinematic analyses and figure plotting. Bova_Leventhal_code.zip contains custom MATLAB software for reconstruction of 3D reach trajectories, processing reach-to-grasp kinematics, and creating the figures in this manuscript using data in the other Extended Data files. Download Extended Data 1, ZIP file.
Extended Data 2
eNeuro_summary_data.zip contains .mat files used by the code in Bova_Leventhal_code.zip to create all main and Extended Data figures, except for Extended Data Figures 6-3–6-16B–D. Download Extended Data 2, ZIP file.
Extended Data Figure 2-1
Individual rat data for hand trajectory variability, reach duration, and reach velocity. A, Average hand trajectory variability of the reach component for learners (left) and non-learners (right) represented as the mean distance from the average trajectory (mm). B, Average hand trajectory variability of the grasp component for learners (left) and non-learners (right) represented as the mean distance from the average trajectory (mm). C, Average reach duration (ms) for learners (left) and non-learners (right). D, Maximum reach velocity (mm/s) for learners (left) and non-learners (right). Data and code to generate this figure are contained in Extended Data 1, 2. Download Figure 2-1, TIF file.
Extended Data Figure 3-1
Individual rat data for reach endpoints. Average reach endpoints of the hand (left) and digit 2 (right) for learners (left columns) and non-learners (right columns) in the X, Y, and Z directions. Pellet is at (0,0,0). Data and code to generate this figure are contained in Extended Data 1, 2. Download Figure 3-1, TIF file.
Extended Data Figure 3-2
Individual rat data for endpoint variability. Average determinant of the covariance matrix (generalized variance) of reach endpoints for the hand and digits. For digits, left two columns show endpoint variability of “raw” digit positions and right two columns show endpoint variability of digit positions subtracted from the hand position. Data and code to generate this figure are contained in Extended Data 1, 2. Download Figure 3-2, TIF file.
Extended Data Figure 4-1
Individual rat fine digit kinematics data. A, Average digit 2 trajectory variability of the reach component for individual rats represented as the mean distance from the average trajectory. B, Average digit 2 trajectory variability of the grasp component for individual rats represented as the mean distance from the average trajectory. C, Average aperture at reach end (mm) for individual rats. D, Average aperture variance at reach end for individual rats. E, Average hand orientation (degrees) at reach end for individual rats. F, Average hand orientation MRL at reach end for individual rats. G, Average digit flexion (degrees) at reach end for individual rats. H, Average digit flexion MRL at reach end for individual rats. Data and code to generate this figure are contained in Extended Data 1, 2. Download Figure 4-1, TIF file.
Extended Data Figure 5-1
Digit and forelimb coordination individual rat data. A, Average grasp aperture at the zdigit2 coordinate (+3 mm) indicated by the dashed line in Figure 5A as a function of day number for individual rats (left learners; right non-learners). B, Average hand orientation (degrees) at the zdigit2 coordinate (+3 mm) indicated by the dashed line in Figure 5C as a function of day number for individual rats. C, Average digit flexion (degrees) at the zdigit2 coordinate (+3 mm) indicated by the dashed line in Figure 5E as a function of day number for individual rats. Data and code to generate this figure are contained in Extended Data 1, 2. Download Figure 5-1, TIF file.
Extended Data Figure 6-1
Trajectory variability by outcome for individual rats. A, Average hand trajectory variability of the reach component for successful reaches (“hits,” left column) and unsuccessful reaches (“misses,” right column) for individual learner (left) and non-learner (right) rats. B, Average hand trajectory variability of the grasp component for successful reaches (“hits,” left column) and unsuccessful reaches (“misses,” right column) for individual learner (left) and non-learner (right) rats. Data and code to generate this figure are contained in Extended Data 1, 2. Download Figure 6-1, TIF file.
Extended Data Figure 6-2
Reach endpoint by outcome for individual rats. A, Average reach endpoint (X direction) for successful reaches (“hits”, left column) and unsuccessful reaches (“misses”, right column) for individual learner (left) and non-learner (right) rats. B, Average reach endpoint (Y direction) for successful reaches (“hits”, left column) and unsuccessful reaches (“misses”, right column) for individual learner (left) and non-learner (right) rats. C, Average reach endpoint (Z direction) for successful reaches (“hits”, left column) and unsuccessful reaches (“misses”, right column) for individual learner (left) and non-learner (right) rats. D, Average reach endpoint variability of the hand for successful reaches (“hits”, left column) and unsuccessful reaches (“misses”, right column) for individual learner (left) and non-learner (right) rats. E, Average reach endpoint variability of digit 2 for successful reaches (“hits,” left column) and unsuccessful reaches (“misses,” right column) for individual learner (left) and non-learner (right) rats. Data and code to generate this figure are contained in Extended Data 1, 2. Download Figure 6-2, TIF file.
Extended Data Figure 6-3
Kinematics summary sheet for a nonlearner rat. A, Success rate across days. Pink lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Pink dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 3. Download Figure 6-3, TIF file.
Extended Data Figure 6-4
Kinematics summary sheet for a learner rat. A, Success rate across days. Green lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Green dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 3. Download Figure 6-4, TIF file.
Extended Data Figure 6-5
Kinematics summary sheet for a nonlearner rat. A, Success rate across days. Pink lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Pink dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 4. Download Figure 6-5, TIF file.
Extended Data Figure 6-6
Kinematics summary sheet for a nonlearner rat. A, Success rate across days. Pink lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Pink dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 4. Download Figure 6-6, TIF file.
Extended Data Figure 6-7
Kinematics summary sheet for a nonlearner rat. A, Success rate across days. Pink lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Pink dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 5. Download Figure 6-7, TIF file.
Extended Data Figure 6-8
Kinematics summary sheet for a learner rat. A, Success rate across days. Green lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Green dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 5. Download Figure 6-8, TIF file.
Extended Data Figure 6-9
Kinematics summary sheet for a nonlearner rat. A, Success rate across days. Pink lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Pink dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 6. Download Figure 6-9, TIF file.
Extended Data Figure 6-10
Kinematics summary sheet for a nonlearner rat. A, Success rate across days. Pink lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Pink dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 7. Download Figure 6-10, TIF file.
Extended Data Figure 6-11
Kinematics summary sheet for a learner rat. A, Success rate across days. Green lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Green dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 8. Download Figure 6-11, TIF file.
Extended Data Figure 6-12
Kinematics summary sheet for a nonlearner rat. A, Success rate across days. Pink lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. A, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Pink dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 9. Download Figure 6-12, TIF file.
Extended Data Figure 6-13
Kinematics summary sheet for a learner rat. A, Success rate across days. Green lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Green dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 10. Download Figure 6-13, TIF file.
Extended Data Figure 6-14
Kinematics summary sheet for a nonlearner rat. A, Success rate across days. Pink lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Pink dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 11. Download Figure 6-14, TIF file.
Extended Data Figure 6-15
Kinematics summary sheet for a nonlearner rat. A, Success rate across days. Pink lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Pink dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 12. Download Figure 6-15, TIF file.
Extended Data Figure 6-16
Kinematics summary sheet for a nonlearner rat. A, Success rate across days. Pink lines indicate first reach success, black lines indicate success on any reach attempt for a single trial. B, Shared control plots illustrating measures of hand shaping at reach end (all data for a single rat that went into Fig. 4C–N). The top axes are swarm plots showing aperture at reach end for every trial. Pink dots indicate first-reach success trials, black dots indicate first reach failed trials (i.e., pellet remained, pellet knocked off, or multiple reach success), and gray dots indicate all other trials (e.g., no pellet delivered). Bottom plots show the difference between the mean value on each day and the mean value on day 1. Distributions show the results of a bootstrap resampling procedure with 95% confidence intervals indicated by the solid lines at the left of each distribution. C, Same as B for the mean distance from the average reach trajectory for each day (top panel shows all hand location data for a single rat that went into Fig. 2C, left panel; bottom panel shows all digit 2 location data that went into Fig. 4A, left panel). D, Same as B, C for the reach endpoint analyses. Left column shows reach endpoints in x, y, and z for the hand location (all data for a single rat that went into Fig. 3B, left panels). Right column shows reach endpoints for digit 2 with respect to the hand location (all data for a single rat that went into Fig. 3B, right panels, except here the hand location was subtracted out). Data and code to generate this figure are contained in Extended Data 1, 2, 13. Download Figure 6-16, TIF file.
Extended Data 3
eNeuro_session_summaries_01.zip contains .mat files needed to create Extended Data Figures 6-3, 6-4B–D. Download Extended Data 3, ZIP file.
Extended Data 4
eNeuro_session_summaries_02.zip contains .mat files needed to create Extended Data Figures 6-5, 6-6B–D. Download Extended Data 4, ZIP file.
Extended Data 5
eNeuro_session_summaries_03.zip contains .mat files needed to create Extended Data Figures 6-7, 6-8B–D. Download Extended Data 5, ZIP file.
Extended Data 6
eNeuro_session_summaries_04.zip contains .mat files needed to create Extended Data Figure 6-9B–D. Download Extended Data 6, ZIP file.
Extended Data 7
eNeuro_session_summaries_05.zip contains .mat files needed to create Extended Data Figure 6-10B–D. Download Extended Data 7, ZIP file.
Extended Data 8
eNeuro_session_summaries_06.zip contains .mat files needed to create Extended Data Figure 6-11B–D. Download Extended Data 8, ZIP file.
Extended Data 9
eNeuro_session_summaries_07.zip contains .mat files needed to create Extended Data Figure 6-12B–D. Download Extended Data 9, ZIP file.
Extended Data 10
eNeuro_session_summaries_08.zip contains .mat files needed to create Extended Data Figure 6-13B–D. Download Extended Data 10, ZIP file.
Extended Data 11
eNeuro_session_summaries_09.zip contains .mat files needed to create Extended Data Figure 6-14B–D. Download Extended Data 11, ZIP file.
Extended Data 12
eNeuro_session_summaries_10.zip contains .mat files needed to create Extended Data Figure 6-15B–D. Download Extended Data 12, ZIP file.
Extended Data 13
eNeuro_session_summaries_11.zip contains .mat files needed to create Extended Data Figure 6-16B–D. Download Extended Data 13, ZIP file.
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