Двойственная природа зеркальной нейронной системы человека: фасилитация и ингибирование при различных протоколах предъявления и стимуляции тема диссертации и автореферата по ВАК РФ 00.00.00, кандидат наук Нието Доваль Карлос Муриель

• Нието Доваль Карлос Муриель
• кандидат наук
• 2025

Оглавление диссертации кандидат наук Нието Доваль Карлос Муриель

TABLE OF CONTENTS

LIST OF ABBREVIATIONS

INTRODUCTION

Relevance of the study

Purpose and objectives of the study

Object and subject of the study

Scientific novelty

Theoretical Contributions

Clinical Relevance

Methodological Innovations

Approbation of scientific results and publications

Main provisions submitted for defense

CHAPTER 1: LITERATURE REVIEW

1.1 Mirror Neurons and Motor Cortex Dynamics

1.2 Transcranial Magnetic Stimulation (TMS) in Human Mirror Neuron Research

1.2.1 Non-Invasive Brain Stimulation (NIBS): General Characteristics and Applications

1.2.2 Transcranial Magnetic Stimulation (TMS)

1.2.3 TMS Implemented with Stimuli Presentation for the Study ofMN

1.3 Motor Inhibition in Motor Control

1.4 Protocols to Study Excitatory and Inhibitory Mechanisms of the Motor Cortex

1.5 Influence of Body Posture in Neural Processes

CHAPTER 2: METHODS

2.1 General Methodological Approach

2.2 Experimental Details (Based on Studies)

2.2.1 Study 1 (Methodological Basis for Stimuli Presentation and TMS Timing)

2.2.2 Study 2 (Temporal Dynamics)

2.2.3 Study 3 (Excitatory/Inhibitory Components)

2.2.4 Study 4 (Body Posture and Mirror Neurons)

CHAPTER 3: RESULTS

3.1 Summary of Key Findings

3.1.1 Study 1 (Methodological Basis for Stimuli Presentation and TMS Timing)

3.1.2 Study 2 (Temporal Dynamics)

3.1.3 Study 3 (Excitatory/Inhibitory Components)

3.1.4 Study 4 (Body Posture and Mirror Neurons)

3.2 Results for Each Study

3.2.1 Study 1 (Methodological Basis for Stimuli Presentation and TMS Timing)

3.2.2 Study 2 (Temporal Dynamics)

3.2.3 Study 3 (Excitatory/Inhibitory Components)

3.2.4 Study 4 (Body Posture and Mirror Neurons)

CHAPTER 4: DISCUSSION

4.1 Study 1 (Methodological Basis for Stimuli Presentation and TMS Timing)

4.1.1 Interpretation of Findings

4.1.2 Implications for Theory and Practice...............................................................Ill

4.1.3 Limitations

4.1.4 Future Directions

4.2 Study 2 (Temporal Dynamics)

4.2.1 Interpretation of Findings

4.2.2 Implications for Theory and Practice

4.2.3 Limitations

4.2.4 Future Directions

4.3 Study 3 (Excitatory/Inhibitory Components)

4.3.1 Interpretation of Findings

4.3.2 Implications for Theory and Practice

4.3.3 Limitations

4.3.4 Future Directions

4.4 Study 4 (Body Posture and Mirror Neurons)

4.4.1 Interpretation of Findings

4.4.2 Implications for Theory and Practice

4.4.3 Limitations

4.4.4 Future Directions

CONCLUSIONS

Summary of Contributions

Final Remarks

REFERENCES

APPENDIX - RUSSIAN TRANSLATION OF THE DISSERTATION

Введение диссертации (часть автореферата) на тему «Двойственная природа зеркальной нейронной системы человека: фасилитация и ингибирование при различных протоколах предъявления и стимуляции»

Introduction

The discovery of mirror neurons (MNs) in the early 1990s within the ventral premotor cortex (area F5) of macaques marked a transformative development in neuroscience, illuminating the neural basis of action understanding and social cognition [1]. Mirror neurons, characterized by their unique activation during both the execution and observation of actions, have since been identified in humans, where they are distributed across regions including the inferior frontal gyrus, precentral gyrus, and inferior parietal lobule [2; 3]. These neurons serve as a crucial interface between perception and motor systems, enabling the internal simulation of observed actions. This functionality underpins various cognitive processes, such as imitation, empathy, and language development [4; 5].

Research into the mirror neuron system (MNS) has revealed its somatotopically organized activation patterns, which correspond to specific body parts or actions, and its differential responsiveness to transitive and intransitive actions [6]. While the MNS autonomously engages during action observation and execution, its functionality can be modulated by contextual and intrinsic factors, such as the type of stimuli presented or neurological conditions [7]. Furthermore, transcranial magnetic stimulation (TMS) has emerged as a powerful tool to investigate the causal mechanisms underlying MNS activity, enabling precise temporal and spatial examination of motor-evoked potentials (MEPs) as proxies for neural responses [8].

Relevance of the study

Despite these advances, significant gaps remain in understanding the intricate balance between excitatory and inhibitory processes within the MNS and the temporal dynamics of its activation. For instance, while motor surround inhibition (mSI) has been

well-documented during action execution, its role during action observation—a hallmark of mirror neuron activity—remains underexplored [9; 10]. Moreover, the influence of contextual factors such as body posture and head orientation on MNS functionality has received limited attention, despite their potential implications for embodied cognition and motor learning [11].

This thesis seeks to address these knowledge gaps by examining the temporal dynamics, excitatory and inhibitory components, and contextual modulations of MNS activity using TMS in a series of four interrelated studies. By leveraging advanced experimental paradigms, such as the novel "Postvideo" condition and paired-pulse TMS protocols, this research aims to elucidate the mechanisms that govern MNS responses and their broader implications for neuroscience and clinical applications.

Purpose and objectives of the study

The overarching aim of this thesis is to deepen the understanding of the MNS by addressing the following key research questions:

1. Optimal Conditions for MNS Activation:

o What is the most effective combination of stimuli presentation and TMS timing for eliciting robust and temporally stable MNS responses?

2. Temporal Dynamics of Mirror Neuron Activity:

o How do MNS responses evolve over time, particularly in terms of excitatory and inhibitory processes, during the observation of goal-directed movements?

3. Excitatory and Inhibitory Components:

o What are the distinct roles of excitatory and inhibitory mechanisms in shaping MNS functionality, and how do these processes interact to modulate motor excitability?

4. Contextual Modulations:

o How do body posture and head orientation influence MNS activity, and what are the implications of these effects for motor learning and rehabilitation?

Object and subject of the study

To address these questions, the research objectives include:

• Establishing a methodological framework for optimizing stimuli presentation and TMS timing to enhance the reliability of MNS studies.

Investigating the timeline of MNS activation, with a particular focus on the inhibitory component (e.g., mSI) as a key modulator of motor responses.

To elucidate the neural mechanisms underlying the excitatory and inhibitory interplay within the MNS using paired-pulse TMS paradigms.

Exploring the influence of body posture and proprioceptive cues on MNS functionality, providing insights into its adaptability and robustness.

Scientific novelty

The findings of this thesis have far-reaching implications for both theoretical neuroscience and clinical practice. By advancing our understanding of the temporal and mechanistic properties of the MNS, this research contributes to foundational knowledge in motor control, social cognition, and neural plasticity. Specifically:

Theoretical Contributions

o The demonstration of temporally specific excitatory and inhibitory patterns enriches models of neural simulation, offering a nuanced perspective on how the MNS encodes and differentiates observed actions.

o The integration of mSI into the framework of MNS activity highlights the parallels between action execution and observation, supporting theories of embodied cognition.

Clinical Relevance

o The identification of optimal conditions for eliciting MNS responses provides a basis for developing targeted interventions in neurorehabilitation. For instance, leveraging the Postvideo condition could enhance motor recovery in patients with stroke or movement disorders by engaging the MNS more effectively.

o Understanding the influence of body posture and proprioceptive states on MNS activity could inform therapeutic strategies for conditions such as dystonia and Parkinson's disease, where motor control and body awareness are disrupted.

Methodological Innovations

o The novel experimental paradigms introduced in this thesis, including the Postvideo condition and paired-pulse TMS protocols, offer robust and replicable tools for future research on MNS dynamics and beyond.

By addressing critical gaps in MNS research and introducing innovative methodologies, this thesis lays the groundwork for future investigations into the neural

mechanisms of action observation and their translational potential. The insights gained extend beyond neuroscience, offering implications for robotics, artificial intelligence, and the design of adaptive learning systems that emulate human motor cognition.

Approbation of scientific results and publications

This thesis was prepared at the Institute for Cognitive Neurosciences, National Research University Higher School of Economics. Three published articles were selected for the

defense procedure:

1. Нието-Доваль К., Рагимова А. А., Феурра M.. (2023). ВЛИЯНИЕ ЗРИТЕЛЬНОГО ПРЕДЪЯВЛЕНИЯ ДВИЖЕНИЙ ПАЛЬЦЕВ РУКИ НА МОТОРНЫЙ ОТВЕТ, ВЫЗВАННЫЙ ТРАНСКРАНИАЛЬНОЙ МАГНИТНОЙ СТИМУЛЯЦИЕЙ. ЭФФЕКТ, СВЯЗАННЫЙ С ВОЗМОЖНОЙ РЕАКЦИЕЙ СИСТЕМЫ ЗЕРКАЛЬНЫХ НЕЙРОНОВ. Журнал высшей нервной деятельности им. И.П. Павлова, 2023, Т. 73, № 3, стр. 334-347 (Russian Version) //Nieto-Doval, С., Ragimova, А. А., & Feurra, М. (2023). Influence of Visual Presentation of Finger Movements on Motor Responses Induced by Transcranial Magnetic Stimulation. An Effect Linked with a Possible Reaction of the Mirror Neuron System. Neuroscience and Behavioral Physiology, 53(8), 1426-1434. https://doi.org/10.1007/sllQ55-023-01535-0 (English Version)

2. Ragimova, A., Doval, C. N., Salamatin, M., Vorobyova, A., Shevtsov, O., Viazmin, A., & Feurra, M. (2024). The Influence of Body Posture on the Mirror Neuron System. Psychology. Journal of Higher School of Economics, 21(3), 439-455.

3. Нието-Доваль К., Рагимова А.А., Вязьмин A.O., Шевцов О.И., Феурра М. Временной профиль активации системы зеркальных нейронов при наблюдении за движением. Исследование при помощи транскраниальной магнитной стимуляции. Нервно-мышечные болезни. 2024;14(4):40-50. https://doi.org/10.17650/2222-8721-2024-14-4-40-5Q

4. Nieto-Doval C., Ragimova A., Perevoznyuk G., et al. Motor resonance and inhibitory mechanisms in action observation as revealed by corticospinal excitability II Scientific Reports. 2025. Vol. 15. P. 23245. https://doi.org/10.1038/s41598-Q25-

03989-3

Main provisions submitted for defense

The thesis presents the following key provisions for defense, each supported by empirical evidence and theoretical analysis derived from the four interconnected studies. These provisions collectively highlight the novel contributions of the research to the field of neuroscience, particularly in understanding the mirror neuron system (MNS) and its role in action observation, motor control, and cognitive processes.

1. Novel Paradigm for Studying Mirror Neuron Dynamics

The introduction of the Postvideo condition, which involves delayed TMS application following the completion of observed movements, provides a robust and temporally stable approach for investigating MNS activity.

This paradigm surpasses traditional methods (Photo and Video conditions) in revealing the dynamic interplay between excitatory and inhibitory components, enabling precise temporal mapping of mirror neuron responses.

2. Identification of a Dual Mechanism in Mirror Neuron Activity

Mirror neuron activation is characterized by a dual mechanism

comprising:

o Excitatory Activation: Enhanced motor-evoked potentials (MEPs) in muscles directly related to the observed movement.

o Inhibitory Suppression: Motor surround inhibition (mSI) in non-related muscles, emphasizing the role of inhibition in maintaining motor precision and preventing unnecessary movements during action observation.

3. Temporal Specificity of Mirror Responses

The research demonstrates that mirror neuron activity follows a distinct temporal trajectory, with the strongest responses observed at later time points (e.g., 640 ms post-movement).

This temporal pattern highlights the significance of the post-movement phase for optimal mirror neuron activation, providing insights into the timing of excitatory and inhibitory processes.

4. Role of Motor Surround Inhibition (mSI)

mSI emerges as a central feature of mirror neuron activity, reflecting a neural mechanism for selectively suppressing non-related muscles while enhancing task-related muscle activation.

• The findings extend the concept of mSI from action execution to action observation, demonstrating its critical role in simulating precise motor control.

5. Independence of MNS from Observer's Body Posture

The study establishes that mirror neuron responses are not significantly influenced by head orientation or neck position, underscoring the system's adaptability across various body postures.

This independence suggests that the MNS primarily relies on visual and cognitive cues, enhancing its role in social cognition and action understanding irrespective of the observer's proprioceptive state.

6. Interaction Between Excitatory and Inhibitory Mechanisms

The correlation between TMS protocols (SICI, ICF, SP) and MEPs during the mirror task highlights the complementary roles of excitation and inhibition in modulating motor responses during action observation.

• The findings provide evidence for the involvement of GABAergic circuits in mirror neuron activity, emphasizing the balance of excitatory and inhibitory mechanisms in motor control.

7. Integration with Cortico-Basal Ganglia Networks

The study suggests a potential interaction between the MNS and the cortico-basal ganglia network, emphasizing the role of inhibitory circuits in fine-tuning motor responses during action observation.

• This interaction extends the understanding of mirror neuron activity within a broader neural framework, linking it to established motor control mechanisms.

8. Practical Implications for Neurorehabilitation

The findings offer a foundation for developing action observation-based therapies targeting motor recovery in patients with stroke, neurodegenerative diseases, or motor impairments.

• The emphasis on inhibitory dynamics and temporal precision in MNS activity provides actionable insights for designing targeted interventions that leverage mirror neuron responses.

These provisions for defense encapsulate the core contributions and implications of the thesis, offering a comprehensive framework for understanding the mirror neuron system's dynamic and multifaceted nature. They underscore the theoretical advancements and practical applications of the research, solidifying its significance in neuroscience and related fields.

Заключение диссертации по теме «Другие cпециальности», Нието Доваль Карлос Муриель

Заключение

Проведенное диссертационное исследование существенно расширяет современные представления о СЗН, раскрывая сложные паттерны взаимодействия процессов возбуждения, торможения и временной динамики во время НД. Разработанная парадигма «Поствидео» доказала свою эффективность как инструмент для комплексного анализа реакций СЗН, демонстрируя их зависимость от временных параметров и специфики двигательной задачи.

Выявленная временная динамика эффектов свидетельствует о максимальном вовлечении зеркальных нейронов на заключительных этапах НД, особенно в момент завершения действия или непосредственно сразу после него. Обнаруженная поздняя активация согласуется с гипотезой о роли СЗН в оценке и симуляции наблюдаемых действий, что вносит существенный вклад в понимание их участия в процессах моторного научения, координации и социального познания. Кроме того, пролонгированный характер JIT подтверждает его функциональную значимость, указывая на ключевую роль тормозных процессов в обеспечении точного и контекстно-зависимого моторного контроля.

Выявление роли JIT как доминирующего компонента реакции СЗН представляет собой концептуально важный результат, противоречащий традиционным представлениям, акцентирующим внимание преимущественно на возбудительных процессах. Полученные данные подчеркивают способность СЗН к селективному подавлению нерелевантных мышц, обеспечивая точное моторное моделирование и минимизируя избыточную активацию. Эти результаты углубляют понимание нейронных механизмов моторного подражания, открывая новые направления для изучения взаимодействия возбуждения и торможения в двигательном познании.

Исследование также подчеркивает высокую адаптационную способность СЗН, что проявляется в ее независимости от проприоцептивных сигналов, включая положение головы. Такая гибкость обеспечивает надежность процессов НД в различных постуральных условиях, подчеркивая эволюционную и функциональную значимость данной системы.

Помимо теоретической значимости, настоящее исследование имеет важное практическое значение для нейрореабилитации. Установленные временные и пространственные характеристики активности СЗН закладывают основу для разработки терапевтических вмешательств, направленных на восстановление моторных функций при таких состояниях, как травмы кисти [154], а также на улучшение двигательной активности у пациентов с неврологическими нарушениями, включая постинсультные состояния [155; 156] и болезнь Паркинсона [157]. Кроме того, акцент на тормозных механизмах открывает новые стратегические подходы к повышению точности и координации движений, что имеет важное значение для развития методов клинической нейрореабилитации.

Изучение нейронных цепей, обеспечивающих функционирование СЗН,

включая их взаимодействие с БГ, мозжечком и другими моторными областями,

является перспективным направлением дальнейших исследований. Особый

интерес представляет применение полученных результатов в клинических

исследованиях, особенно в контексте нейропластичности и моторного обучения.

Потенциал использования динамических характеристик СЗН в терапевтических

289

вмешательствах представляет собой перспективное направление нейронаучных исследований, открывающее новые возможности для интеграции фундаментальных знаний и практических применений.

В заключение следует отметить, что настоящее диссертационное исследование формирует прочную теоретическую и методологическую основу для понимания СЗН, подчеркивая сложную организацию ее динамических характеристик и функциональную значимость. Полученные результаты создают надежную базу для развития теоретических моделей, практических приложений и дальнейших исследований в области двигательного познания и смежных дисциплинах.

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