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Research ArticleResearch Article: New Research, Cognition and Behavior

Dynamic Encoding of Reward Prediction Error Signals in the Pigeon Ventral Tegmental Area during Reinforcement Learning

Zhigang Shang, Jiashuo Zhang, Mengmeng Li, Suchen Li, Yinghui Wang and Lifang Yang
eNeuro 19 February 2026, 13 (3) ENEURO.0355-25.2026; https://doi.org/10.1523/ENEURO.0355-25.2026
Zhigang Shang
1School of Electrical and Information Engineering, Zhengzhou University, Zhengzhou 450001, China
2Henan Key Laboratory of Brain Science and Brain-Computer Interface Technology, Zhengzhou 450001, China
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Jiashuo Zhang
1School of Electrical and Information Engineering, Zhengzhou University, Zhengzhou 450001, China
2Henan Key Laboratory of Brain Science and Brain-Computer Interface Technology, Zhengzhou 450001, China
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Mengmeng Li
1School of Electrical and Information Engineering, Zhengzhou University, Zhengzhou 450001, China
2Henan Key Laboratory of Brain Science and Brain-Computer Interface Technology, Zhengzhou 450001, China
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  • ORCID record for Mengmeng Li
Suchen Li
1School of Electrical and Information Engineering, Zhengzhou University, Zhengzhou 450001, China
2Henan Key Laboratory of Brain Science and Brain-Computer Interface Technology, Zhengzhou 450001, China
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Yinghui Wang
1School of Electrical and Information Engineering, Zhengzhou University, Zhengzhou 450001, China
2Henan Key Laboratory of Brain Science and Brain-Computer Interface Technology, Zhengzhou 450001, China
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Lifang Yang
1School of Electrical and Information Engineering, Zhengzhou University, Zhengzhou 450001, China
2Henan Key Laboratory of Brain Science and Brain-Computer Interface Technology, Zhengzhou 450001, China
3The Affiliated Encephalopathy Hospital of Zhengzhou University, Zhumadian 463000, China
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  • Figure 1.
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    Figure 1.

    Task design, recording setup, and VTA electrode localization. a, Schematic of the reinforcement learning experimental paradigm. In each trial, either a green (cue+) or red (cue−) LED light is randomly illuminated for 2 s. If the pigeon pecks the green key during this period, it receives a 2 s food reward, followed by a 2 s intertrial interval. Pecks on the red key or no response result in no reward and immediate transition to the next trial. For neural analyses, precue/cue/outcome epochs were defined as 0.5 s windows aligned to cue onset and outcome time. b, A diagram of the experimental apparatus. c, Pigeon with implanted electrode. d, Electrode implantation site, histological verification, raw traces and detected MUA spike events from the 16 recording channels.

  • Figure 2.
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    Figure 2.

    Behavioral learning curves and representative VTA spike waveforms. a, Key pecking accuracy of P109, P117, and P121 across sessions. b, Spike waveform samples after preprocessing.

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    Figure 3.

    Learning-related redistribution of VTA pooled MUA across task epochs during cue+ trials. a, Example cue+ rasters (left) and PSTHs (right) for pigeon P109 across sessions, aligned to cue onset (0.5 s) and reward delivery (1.0 s; dashed lines). Background shading indicates sessions assigned to the learning phase (blue) versus the consolidation phase (pink) based on behavioral performance. For each session, PSTHs are shown at two temporal resolutions (upper, 10 ms bins; lower, 100 ms bins with a 5-point moving average for visualization). b, Session-wise spike proportions for cue+ trials in the precue reference, cue+, and reward epochs for each pigeon (mean ± SD across valid trials). Statistical significance is indicated above bars (see Materials and Methods, Statistical analysis). c, Representative example from pigeon P109 illustrating how the session-wise mean spike proportion evolves across training. Each point corresponds to the same mean value shown for P109 in panel b (computed across valid trials within that session for the cue and reward epochs). The curve is provided as a visualization of the learning-related trajectory rather than a group-level summary. d, Session-to-session change in the same metric for P109, computed as Δ = value (Session n + 1) − value (Session n), where each value is the session-wise mean spike proportion (identical to the corresponding bar height in panel b).

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    Figure 4.

    Cue− trials show limited cue-locked modulation in VTA pooled MUA. a, Example cue− rasters (left) and PSTHs (right) for pigeon P109 across Sessions 1–3, aligned to cue− onset (dashed line). The y-axis indexes valid trials after trial-level quality control. For each session, PSTHs are shown at two temporal resolutions (upper, 10 ms bins; lower, 100 ms bins with a 5-point moving average for visualization). b, Session-wise comparison of cue− spike proportions between the precue reference and cue− epochs for each pigeon (mean ± SD across valid trials). “n.s.” indicates no significant difference between epochs (see Materials and Methods, Statistical analysis).

Tables

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    Table 1.

    Summary of trial inclusion/exclusion and recording channels (pooled MUA) by pigeon

    Pigeon ID (sex)Total trials/valid trials/ excluded trialsExclusion reasons (trial count)Number of channels
    P109 (female)267/210/57Premature response (<500 ms; n = 35) large-amplitude wing flapping (n = 22)16
    P117 (female)153/89/64Premature response (<500 ms; n = 33) large-amplitude wing flapping (n = 31)16
    P121 (male)319/152/167Premature response (<500 ms; n = 97) large-amplitude wing flapping (n = 70)16
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    Table 2.

    Session-wise summary of trial counts used for pooled MUA analyses

    Pigeon IDSessionExtracted key-peck trialsValid trialsChannels included (n)
    P109S1412816
    S2553916
    S3534016
    S4615416
    S5574916
    P117S1452116
    S2573516
    S3513316
    P121S1632616
    S2723316
    S3683416
    S4673216
    S5492716
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eneuro: 13 (3)
eNeuro
Vol. 13, Issue 3
March 2026
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Dynamic Encoding of Reward Prediction Error Signals in the Pigeon Ventral Tegmental Area during Reinforcement Learning
Zhigang Shang, Jiashuo Zhang, Mengmeng Li, Suchen Li, Yinghui Wang, Lifang Yang
eNeuro 19 February 2026, 13 (3) ENEURO.0355-25.2026; DOI: 10.1523/ENEURO.0355-25.2026

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Dynamic Encoding of Reward Prediction Error Signals in the Pigeon Ventral Tegmental Area during Reinforcement Learning
Zhigang Shang, Jiashuo Zhang, Mengmeng Li, Suchen Li, Yinghui Wang, Lifang Yang
eNeuro 19 February 2026, 13 (3) ENEURO.0355-25.2026; DOI: 10.1523/ENEURO.0355-25.2026
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Keywords

  • dynamic encoding
  • pigeon
  • reward prediction error
  • spike
  • ventral tegmental area

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