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Premotor cortex and the recognition of motor actions

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Abstract

In area F5 of the monkey premotor cortex there are neurons that discharge both when the monkey performs an action and when he observes a similar action made by another monkey or by the experimenter. We report here some of the properties of these 'mirror' neurons and we propose that their activity 'represents' the observed action. We posit, then, that this motor representation is at the basis of the understanding of motor events. Finally, on the basis of some recent data showing that, in man, the observation of motor actions activate the posterior part of inferior frontal gyrus, we suggest that the development of the lateral verbal communication system in man derives from a more ancient communication system based on recognition of hand and face gestures.

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... The mirror neurons were found in the premotor cortex of apes (Di Pellegrino et al., 1992;Gallese et al., 1996). The anatomical homology between the area F5 and the Broca's area led to the hypothesis that the neurons with functional characteristics of mirror neurons exist in the human cortex.The functionality of these cells in displaying through the visual observation of an action or through the perception of a hearing stimulus Kohler et al., (2002) suggests that this class of neurons computes the meaning of an action or of a sound based on the repertoire of sensory-motor schemes of the observer/receptor (Gallese, 2003;Rizzolatti et al., 1996). Noting that the ideomotor apraxia neurocognitive disturb is similar to the functional characteristics of the mirror neurons from the area F5, speculates that this class of neural cells computes the production and perception of phonetic gestures in the Broca's area. ...
... The hypothesis that the mirror neurons compute the syntax was substantiated on the assumption that this class of cells operates executive aspects of the speech Rizzolatti et al., (1996); Rizzolatti e Arbib (1998). The sequence of articulated movements to do an activity goes through the motor cortex, which mediates the organization of sentences of action. ...
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Neurolinguistics investigates the structural and functional relationship between the brain and the language. The neurobiological substrates of the natural languages receive, process and transmit linguistic codes which are calculated by the speaker/listener sensory, motor and cognitive systems. We suggest a theoretical reflection about how the understanding of the neuroscience of the language can be interpreted with the language theory of Chomsky from the mirror neurons. We mention and exemplify empirical and theoretical problems based on the assumption that the mirror neurons mediate the computation of the language faculty. We suggest a computational approach as a way of investigating the language faculty from the neurolinguistics perspective. We conclude that the decomposition of the language faculty into elementary components at the neurobiological level and the fragmentation of conceptual totalities can lead it to a hyper reductionism.
... Brain activation during AO has been intensively investigated since the discovery of the firing of "mirror neurons" in area F5 of a monkey's premotor cortex during AO [7][8][9]. These neurons were found to be activated both when the monkey executed an action and when the monkey observed a similar action performed by another monkey or the experimenter [8]. ...
... Brain activation during AO has been intensively investigated since the discovery of the firing of "mirror neurons" in area F5 of a monkey's premotor cortex during AO [7][8][9]. These neurons were found to be activated both when the monkey executed an action and when the monkey observed a similar action performed by another monkey or the experimenter [8]. Consistent with these studies, brain imaging studies in humans have shown that AO induces activity in the motor-related brain areas [10,11]. ...
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Transcranial magnetic stimulation studies have indicated that action observation (AO) modulates corticospinal excitability. Although a few previous studies have shown that the AO of simple motor movements at a slow playback speed facilitates corticospinal excitability more than that at normal playback speed, it is unclear if this effect occurs during the AO of sport-related complex movements. Therefore, we investigated the changes in the motor evoked potential (MEP) amplitudes of the flexor carpi radialis (FCR) and abductor digiti minimi (ADM) muscles during the AO of a basketball free-throw movement at three different playback speeds (100%, 75%, and 50% speeds). Additionally, we evaluated the effects of stimulus timing (holding the ball vs. releasing the ball for shooting) and motor expertise (expert basketball players vs. novices) on the MEP amplitude during the AO. Our results demonstrated that regardless of motor expertise, the MEP amplitude of the FCR muscle was significantly smaller in the 50% speed condition than in the 100% condition. In the ADM muscle, the MEP amplitude was significantly larger when the ball was held after dribbling than when the ball was released. Therefore, it is suggested that corticospinal excitability in specific muscles during the observation of complex whole-body movements is influenced by video playback speed and stimulus timing.
... Током истраживања префронталне функције код супхуманих примата откривени су тзв. огледалски неурони (Rizzolatti et al., 1996) који се налазе у дорзолатералном делу фронталног режња. Они се активирају не само када животиња обавља неку активност него и када посматра другог који обавља исту активност. ...
... For example, one of the functions of imitation in early development is its connection with social skills (Krstić, 2008, p. 60). While examining prefrontal functions in subhuman primates, so-called mirror neurons (Rizzolatti et al., 1996) that are found in the dorsolateral area of the frontal lobe have been discovered. They are activated not only when an animal performs a certain activity, but also when it observes someone else performing the same activity. ...
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Neurosociology is a relatively new interdisciplinary field of science. It is a discipline that lies on the borders between sociology, social psychology, biology and neurosciences. This scientific field offers an opportunity for better understanding interactions between brain and interpersonal functioning by analyzing the influence of social factors on creating brain organization and its functions.
... Girard describes the process by which humans' mimicry leads to rivalry and violence as admiration for a model turns into violent hostility against a rival, and he describes how rivalries between many rivals converge on a single target: a common enemy against whom the group can reunify (Girard, 1972). Corroborated by eminent works in learning theories, development psychology (Meltzoff, 1995, Garrels, 2011 and cognitive neurosciences Rizzolatti et al., 1996), the mimetic theory fertilised cross-disciplinary research and gave birth to a new branch of neuropsychology (Girard et al., 1978, Girard, 2008Oughourlian, 2007Oughourlian, , 2013. The Girardian paradigm offers a promising framework for analysing digital interactions. ...
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We combine philosophical theories with quantitative analyses of online data to propose a sophisticated approach to social media influencers. Identifying influencers as communication systems emerging from a dialectic interactional process between content creators and in-development audiences, we define them mainly using the composition of their audience and the type of publications they use to communicate. To examine these two parameters, we analyse the audiences of 619 Instagram accounts of French, English, and American influencers and 2,400 of their publications in light of Girard’s mimetic theory and McLuhan’s media theory. We observe meaningful differences in influencers’ profiles, typical audiences, and content type across influencers’ classes, supporting the claim that such communication systems are articulated around ‘reading contracts’ upon which influencers’ image is based and from which their influence derives. While the upkeep of their influence relies on them sticking to this contract, we observe that successful influencers shift their content type when growing their audiences and explain the strategies they implement to address this double bind. Different types of contract breaches then lead to distinct outcomes, which we identify by analysing various types of followers’ feedback. In mediating social interactions, digital platforms reshape society in various ways; this interdisciplinary study helps understand how the digitalisation of social influencers affects reciprocity and mimetic behaviours.
... The action observation network in particular includes structures involved in the mirror neuron system (Mizuguchi & Kanosue, 2017;Rizzolatti & Craighero, 2004). Mirror neurons are specialized types of neurons found in the human frontal, parietal and temporo-occipital cortices (Caspers et al., 2010) that fire not only when an individual performs an action, but when they observe actions performed by another (di Pellegrino et al., 1992;Rizzolatti & Craighero, 2004;Rizzolatti et al., 1996). Multiple studies utilizing transcranial magnetic stimulation (TMS) to probe the motor cortex have observed increased corticospinal excitability in response to AO interventions (see Loporto et al., 2011;Naish et al., 2014 for review). ...
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Limb immobilization causes rapid declines in muscle strength and mass. Given the role of the nervous system in immobilization‐induced weakness, targeted interventions may be able to preserve muscle strength, but not mass, and vice versa. The purpose of this study was to assess the effects of two distinct interventions during 1 week of knee joint immobilization on muscle strength (isometric and concentric isokinetic peak torque), mass (bioimpedance spectroscopy and ultrasonography), and neuromuscular function (transcranial magnetic stimulation and interpolated twitch technique). Thirty‐nine healthy, college‐aged adults (21 males, 18 females) were randomized into one of four groups: immobilization only ( n = 9), immobilization + action observation/mental imagery (AOMI) ( n = 10), immobilization + neuromuscular electrical stimulation (NMES) ( n = 12), or control group ( n = 8). The AOMI group performed daily video observation and mental imagery of knee extensions. The NMES group performed twice daily stimulation of the quadriceps femoris. Based on observed effect sizes, it appears that AOMI shows promise as a means of preserving voluntary strength, which may be modulated by neural adaptations. Strength increased from PRE to POST in the AOMI group, with +7.2% (Cohen's d = 1.018) increase in concentric isokinetic peak torque at 30°/s. However, NMES did not preserve muscle mass. Though preliminary, our findings highlight the specific nature of clinical interventions and suggest that muscle strength can be independently targeted during rehabilitation. This study was prospectively registered: ClinicalTrials.gov NCT05072652.
... Another important factor in dynamic intelligence is imitation based on embodiment. The importance of imitation is also emphasized in brain science with mirror neurons [15]. The existence of mirror neurons implies that a human interprets another human's actions as actions by himself, providing an understanding of their meaning. ...
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The agent technology has recently become one of the most vibrant and fastest growing areas in information technology. On of the most promising characteristics of agent is its intelligence. Intelligent agent is the agent that percepts its environment, collects all information about its environment that it needs, processes these information and then generate proper actions according to these information. This paper discusses trends and differences between two main types of intelligence that can be applied to agent: accumulative intelligence and dynamic intelligence. Accumulative intelligence is discussed with its two perspectives: moment perspective and historical perspective. Auto-vehicle driver is also discussed as an application example of accumulative intelligence. Also, MOSAIC, Mimesis, and MINDY models are reviewed as the pioneering works of dynamic intelligence.
... È una dialettica di desiderio, ove il desiderio del bambino è quello di essere desiderato, e tutto quello che lo fa sentire desiderato riuscirà a produrlo all'ennesima potenza, diventando col tempo la base salda per il comportamento adulto. In questo scenario, hanno probabilmente un ruolo importante i neuroni-specchio, gruppi di popolazioni neuronali che inducono a "copiare" i comportamenti di chi abbiamo davanti (Rizzolatti 1996, Leskowitz 2010. Chiaramente a questo stadio, il ridere del bambino è ben altra cosa dal ridere dell'adulto. ...
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The present paper aims at describing the current state of healthcare clown organizations in Latin America, in comparison with organizations in other latitudes. To articulate this paper, a review of online papers available in databases such as WoS, Scopus, and Springer, was used to gather information, with keywords such as healthcare clown and hospital clown in Spanish. Moreover, online research in websites of healthcare clown organizations was included. Analysis indicates that there are still a lot of challenges for professional organizations in the region. Socioeconomic characteristics of Latin America countries translate into challenges for healthcare clown organizations, such as training and fundraising, and more research is urgently needed.
... In social cognition, too, Simulation Theory has been endorsed by the discovery of mirror neurons that enable automatic mimicry at the neural level (Rizzolatti et al., 1996). ...
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Fiction may function to support human social interaction by cultivating empathic abilities. The past decade has yielded promising evidence in support of this theory, though the multidimensionality of both fiction-engagement and empathy have presented methodological challenges and led to mixed findings. Studies have tended to focus on reading and have generally treated cognition as a solely internal process. I position empathy and engagement with fiction as ontologically extended processes. I argue that further systematic exploration of fiction technologies would promote a comprehensive and culturally relevant account of the relationship between fiction-engagement and empathy and enhance understanding of its cognitive architecture.
... Regarding the sense of active grasping, observing mirrored movement increases the activity of the motor network in the brain that is responsible for action observation, cognitive control, attention shift 32 and reorganization of the sensorimotor cortex 33 . In this case, the concept of mirror therapy is triggered, as if one is performing the motion independently 34 . ...
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Accomplishing motor function requires multimodal information, such as visual and haptic feedback, which induces a sense of ownership (SoO) over one’s own body part. In this study, we developed a visual–haptic human machine interface that combines three different types of feedback (visual, haptic, and kinesthetic) in the context of passive hand-grasping motion and aimed to generate SoO over a virtual hand. We tested two conditions, both conditions the three set of feedback were synchronous, the first condition was in-phase, and the second condition was in antiphase. In both conditions, we utilized passive visual feedback (pre-recorded video of a real hand displayed), haptic feedback (balloon inflated and deflated), and kinesthetic feedback (finger movement following the balloon curvature). To quantify the SoO, the participants’ reaction time was measured in response to a sense of threat. We found that most participants had a shorter reaction time under anti-phase condition, indicating that synchronous anti-phase of the multimodal system was better than in-phase condition for inducing a SoO of the virtual hand. We conclude that stronger haptic feedback has a key role in the SoO in accordance with visual information. Because the virtual hand is closing and the high pressure from the balloon against the hand creates the sensation of grasping and closing the hand, it appeared as though the person was closing his/her hand at the perceptual level.
... First discovered in monkey ventral premotor and inferior parietal cortex, mirror neurons (MNs) are a class of visuomotor neurons that activate not only during the execution of a motor act, but also during observation of a similar motor act made by another agent. [19][20][21] Subsequent studies employing brain imaging and electrophysiological techniques have demonstrated that the observation of motor acts activates a similar system in humans. 22,23 This activation is somatotopically organized. ...
Article
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In the past two decades several attempts have been made to promote a correct diagnosis and possible restorative interventions in patients suffering from Disorders of Consciousness (DOC). Sensory stimulation has been proved to be useful in sustaining the level of arousal/awareness and to improve behavioral responsiveness with a significant effect on oro-motor functions. Recently, action observation has been proposed as a stimulation strategy in DOC patients, based on neurophysiological evidence that the motor cortex can be activated not only during action execution, but also when actions are merely observed in the absence of motor output, or during listening to action sounds and speech. This mechanism is provided by the activity of mirror neurons. In the present study, a group of patients with DOC (11 males, 4 females; median age: 55 years; age range 19-74 years) underwent task-based functional MRI in which they had, in one condition, to observe and listen to the sound of mouth actions, in another condition, to listen to verbs with motor or abstract content. In order to verify the presence of residual activation of the mirror neuron system, brain activations of patients were compared with that of a group of healthy individuals (7 males, 8 females; median age: 33.4 years; age range: 24-65 years) preforming the same tasks. The results show that brain activations were lower in DOC patients compared with controls, except for primary auditory areas. During the audiovisual task, 5 out of 15 DOC patients showed only residual activation of low-level visual and auditory areas. Activation of high-level parieto-premotor areas was present in 6 patients. During the listening task, 3 patients showed only low-level activations, and 6 patients activated also high-level areas. Interestingly, in both tasks, one patient with a clinical diagnosis of vegetative state showed activations of high-level areas. Region of interest analysis on Blood Oxygen Level Dependent (BOLD) signal change in temporal, parietal and premotor cortex revealed a significant linear relation with the level of clinical functioning, assessed with Coma Recovery Scale-Revised. We propose a classification of the patient’s response based on the presence of low-level and high-level activations, combined with patient’s functional level. These findings support the use of action observation and listening as possible stimulation strategies in DOC patients and highlight the relevance of combined methods based on functional assessment and brain imaging to provide more detailed neuroanatomical specificity about residual activated areas at both cortical and subcortical levels.
... Ріоццолатті, 1992), і як дзеркало, автоматично відображають чужу поведінку у мозку людини та дозволяють відчути те, що вона бачить, так як якщо б вона робила відповідні дії самі. Тобто коли індивід бачить усміхнену людину мозок дає команду: підняти настрій [7]. Стійкість окремих емоційних проявів спричиняють їх закріплення та виникненню відносно стійких форм поведінки -основи формування характеру. ...
... So, we deny them awareness, we deny them purposeful action, and we know that even if we understand the language they speak to us, they do not understand their language themselves. Where in the presented model is the place for mirror neurons and for the role of Fogassi, Rizzolatti, 1996;. It does not dispute the objections raised by Hickok, Pascolo and Dinstein (Hickok 2009;Pascolo, Ragogna, Rossi, 2009;Dinstein, Thomas, Behrmann, Heeger, 2008). ...
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The hierarchical structure of metaphors, as linguistic concepts and other mind tools embodied in Lakoff's cognitive linguistics terms, requires an indication of the mental representations in human minds and the corresponding biophysical representations. The author claims that the searched biophysical representations, which are the basis of mental representations, may be hierarchically related neurons called semblions. According to the connectionist concept, the hierarchical structure of a neural network emerges during the learning process and concurrently constitutes a memory. Semblions of objects, concepts, ideas, and models can be associated on a neurological level with neural representations of phonemes heard simultaneously while perceiving objects and creating their representations. This way, new higher-order semblions are made, corresponding to the words. The work shows how stimulating semblions can cause recall, association, thinking, and other higher mental functions necessary for using natural language. In this way, the reconstruction of mental states takes place based on the propagation of neuronal excitations. The author also indicates why semblions representing mathematical concepts adequately describe many physical world phenomena. At the same time, their polymodal counterparts can be used for a metaphorical description of qualia. The complementarity of the associative memory model proposed by Horzyk with the model of the architecture of self-conscious systems proposed by Galus, which can be used to build artificial self-conscious systems that reasonably use both natural and formal languages, was pointed out. The further development of recursive models of consciousness leading to the presentation of the Motivated Emotional Mind (MEM) model has confirmed that in the process of bottom-up information transfer responsible for creating semblions, rules of similarity are utilized from engrams representing knowledge in the mind to patterns of neural excitations transmitted from the lower sensory layers of semblions. The functioning of the multilayered architecture of brain fields based on the principles of searching for similarity patterns aligns with the hypothesis of using metaphors in subsequent processes of generalization and association, forming the basis for higher mental functions, and creating the semantics and syntax of the language.
... The concept of covert motor imagery also invokes the related function of action observation. When people or animals observe others performing a motor action, it engages motor-related brain areas in a way similar to self-initiated movement 15,[35][36][37][38][39][40][41][42] . This correlation between observation and action exists even in the activity of individual cortical neurons 12,35,[42][43][44] , which supports the notion of a 'mirror neuron' network through which the motor system can presumably learn new skills by observing others 45 . ...
Article
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The most prominent characteristic of motor cortex is its activation during movement execution, but it is also active when we simply imagine movements in the absence of actual motor output. Despite decades of behavioural and imaging studies, it is unknown how the specific activity patterns and temporal dynamics in motor cortex during covert motor imagery relate to those during motor execution. Here we recorded intracortical activity from the motor cortex of two people who retain some residual wrist function following incomplete spinal cord injury as they performed both actual and imagined isometric wrist extensions. We found that we could decompose the population activity into three orthogonal subspaces, where one was similarly active during both action and imagery, and the others were active only during a single task type—action or imagery. Although they inhabited orthogonal neural dimensions, the action-unique and imagery-unique subspaces contained a strikingly similar set of dynamic features. Our results suggest that during motor imagery, motor cortex maintains the same overall population dynamics as during execution by reorienting the components related to motor output and/or feedback into a unique, output-null imagery subspace.
... Moreover, the concept of affordance, adopted from Gibson's ecological psychology theory [2], has been a widelyused concept in industrial design and product architecture [3][4][5][6][7][8][9][10][11][12]. From a broader perspective, the theory of affordances has been extended and exploited in a diverse range of fields including ecological psychology [13][14][15] [16][17][18][19][20], neuroscience [21][22][23][24][25][26][27][28][29][30], architecture and environmental design [31][32][33][34][35][36][37][38][39][40][41][42][43], human-computer interaction (HCI) [44][45][46][47][48][49][50][51][52], marketing and management [53][54][55][56][57][58][59][60][61][62][63][64][65][66][67], education and pedagogy [68][69][70][71][72], digital work and socialization , and robotics and artificial intelligence (AI) [102][103][104][105][106][107][108][109]. A pictorial overview of the abovementioned fields is presented in Fig. 1. ...
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For centuries, the relationship between form and function has been a point of debate in the communities of architecture and design, leading to the development of various theories which have attempted to establish a tangible relationship between these two entities. Besides, the concept of affordance, adopted from Gibson’s ecological psychology theory, has appeared as a widely-used concept in design practice and research. Nevertheless, while it is generally accepted that these concepts have close dependencies and interactions, it appears that there is no explicit theoretical framework that relates three of the most fundamental concepts of design, namely form, function, and affordance. This paper aims to analyze the concept of affordance in the context of industrial design, where we attempt to develop insights into the role of affordances in relation to form and function. To this end, we define the form-affordance-function (FAF) triangle of design as a major contributor to the establishment of a partial product design specification (PDS) in the design process. We present several examples to investigate the position of affordances in competition with other design considerations such as engineering performance, ergonomics, and aesthetics. The insights into these relationships could have potential implications for designers in making informed early-stage design decisions.
... One crucial issue in Joint Actions is that of visuomotor interference (VMI, Blakemore & Frith, 2005;Kilner et al., 2003). Observing another agent performing a movement is known to activate the corresponding motor plan in the observer (Rizzolatti et al., 1996), influencing their movement execution. An incongruent observed action produces a disruption of the observer's motor plan and, consequently, poorer motor performance (Brass et al., 2001;Cracco et al., 2015;Cracco & Brass, 2019;Forbes & Hamilton, 2017). ...
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Joint actions are defined as coordinated interactions of two or more agents towards a shared goal, often requiring different and complementary individual contributions. However, how humans can successfully act together without the interfering effects of observing incongruent movements is still largely unknown. It has been proposed that interpersonal predictive processes are at play to allow the formation of a Dyadic Motor Plan, encompassing both agents` shares. Yet, direct empirical support for such an integrated motor plan is still limited. In this study, we aimed at testing the properties of these anticipated representations. We collected EEG data while human participants (N = 36; 27 females) drew shapes simultaneously to a virtual partner, in two social contexts: either they had to synchronize and act jointly, or they performed the movements alongside, but independently. We adopted a multivariate approach to show that the social context influenced how the upcoming action of the partner is anticipated during the interval preceding the movement. We found evidence that acting jointly induces an encoding of the partner`s action that is strongly intertwined with the participant`s action, supporting the hypothesis of an integrative motor plan in joint but not in parallel actions.
... So, we deny them awareness, we deny them purposeful action, and we know that even if we understand the language they speak to us, they do not understand their language themselves. Where in the presented model is the place for mirror neurons and for the role of Fogassi, Rizzolatti, 1996;. It does not dispute the objections raised by Hickok, Pascolo and Dinstein (Hickok 2009;Pascolo, Ragogna, Rossi, 2009;Dinstein, Thomas, Behrmann, Heeger, 2008). ...
Article
Full-text available
The hierarchical structure of metaphors, as linguistic concepts and other mind tools embodied in Lakoff's cognitive linguistics terms, requires an indication of the mental representations in human minds and the corresponding biophysical representations. The author claims that the searched biophysical representations, which are the basis of mental representations, may be hierarchically related neurons called semblions. According to the connectionist concept, the hierarchical structure of a neural network arises in learning and is also a memory. Semblions of objects, concepts, ideas, and models can be associated on a neurological level with neural representations of phonemes heard simultaneously while perceiving objects and creating their representations. This way, new higher-order semblions are made, corresponding to the words. The work shows how stimulating semblions can cause recall, association, thinking, and other higher mental functions necessary for using natural language. In this way, the reconstruction of mental states takes place based on the propagation of neuronal excitations. The author also indicates why semblions representing mathematical concepts adequately describe many physical world phenomena. At the same time, their polymodal counterparts can be used for a metaphorical description of qualia. The complementarity of the associative memory model proposed by Horzyk with the model of the architecture of self-conscious systems proposed by Galus, which can be used to build artificial self-conscious systems that reasonably use both natural and formal languages, was pointed out. The further development of recursive models of consciousness leading to the presentation of the Motivated Emotional Mind (MEM) model has confirmed that in the process of bottom-up information transfer responsible for creating semblions, rules of similarity are utilized from engrams representing knowledge in the mind to patterns of neural excitations transmitted from the lower sensory layers of semblions. The functioning of the multilayered architecture of brain fields based on the principles of searching for similarity patterns aligns with the hypothesis of using metaphors in subsequent processes of generalization and association, forming the basis for higher mental functions. The functioning of the multilayered architecture of brain fields based on the principles of searching for similarity patterns aligns with the hypothesis of using metaphors in subsequent processes of generalization and association, forming the basis for higher mental functions, and creating the semantics and syntax of the language.
... Similarly, when we hear music, we parse the elements of the music through, for example, body movement, such as foot tapping or body-sway. At times, our comprehension of the actions responsible for producing music is undetected at a conscious level, but activation of so-called 'mirror neurons' in the brain (e.g., Rizzolatti, Fadiga, Gallese & Fogassi, 1996) reveal its presence nonetheless. ...
Conference Paper
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Body movement displayed in music performance is said to be an overt manifestation of the musician's expressive intentions and goals regarding the music being played. We are interested in studying if different levels of expression result in different amounts of body movement and gestures. For this study, musicians were asked in multiple sessions to play an excerpt from the same piece using three different levels of expression while their movements were recorded using an optical motion capture system. Statistical tests show that an increasing amount of expression resulted in more body movement and that the amount of expression and physical movement were intertwined. Also, we present data suggesting that the head and shoulders travelled a further distance overall, and showed bigger differences between performance manners, compared to the fingers, wrists and lower back. We hypothesize that this is related to the contrasting roles these parts of the body play in piano performance.
... Stemming from the discovery of mirror neurons in macaques (Di Pellegrino et al., 1992;Gallese et al., 1996;Rizzolatti et al., 1996) and the theory of embodied simulation theory that followed (Gallese, 2005(Gallese, , 2014Gallese & Sinigaglia, 2011), a growing body of neuroimaging evidence substantiates the claim that action execution and perception are linked in humans (see Gallese, 2009). As neuroimaging techniques have made it possible to investigate the role of sensorimotor engagement in musical experience, multiple studies demonstrate that motor brain areas are activated during passive music listening (Callan et al., 2006;Chen et al., 2008aChen et al., , 2008bGordon et al., 2018;Halpern & Zatorre, 1999;Halpern et al., 2004;Hickok et al., 2003;Zatorre et al., 1996) as well as during beat entrainment (Grahn & Brett, 2007;Nistri et al., 2006). ...
Article
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Embodied music cognition predicts that our understanding of human-made sounds relates to our experience of making the same or similar movements and sounds, which involves imitation of the source of visual and auditory information. This embodiment of sound may lead to numerous kinetic cross-modal correspondences (CMCs). This article investigates music experience in participants with a non-professionally trained music background across three musical dimensions: Contour (Ascending, Descending, Flat), Vertical Density (Low, Medium, High), and Note Pattern (Binary, Ternary, Quaternary). In order that stimuli should reflect contemporary musical usage yet be subject to a high degree of experimental control, 27 ten-second digital piano tracks were created in collaboration with a film composer. In Study 1, participants were asked to rate the stimuli for perceived Direction, Rotation, Movement, and Emotional and Physical Involvement. We test the effects of these factors in terms of the following theories: general and vocal embodied responses to music, the Ecological Theory of Rotating Sounds, and the Shared Affective Motion Experience model of emotion induction. Results for Study 1 were consistent with theories of general and vocal embodied responses to music, as well as with theories of embodied emotional contagion in music. Study 1 also revealed potential confounds in the stimuli, which were further investigated in Study 2 with a new set of participants rating the stimuli for perceived Pitch, Loudness, and Speed. Results for Study 2 served to dissociate intrinsic features of the stimuli from CMCs. Taken together, the two studies reveal a range of embodied CMCs. Although there are limitations to a perceptual study such as this, these stimuli stand to benefit future research in further investigating the embodiment of musical motion.
... Several studies have reported that action observation induces brain activation corresponding to the production of these movements 25 . In addition to the mirror neurons that had initially been found in premotor cortex 26 , neurons in primary motor cortex M1 also respond during action observation 27,28 . Two functional types of neurons have been found in M1, those that facilitate action execution and those that suppress it 29 . ...
Article
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Which factors influence the perception of our hand location is a matter of current debate. Here, we test if sensorimotor processing contributes to the perception of hand location. We developed a novel visuomotor adaptation procedure to measure whether actively performing hand movements or passively observing them, influences visual perception of hand location. Participants had to point with a handheld controller to a briefly presented visual target. When they reached the remembered position of the target, the controller presented a tactile buzz. In adaptation trials, the tactile buzz was presented when the hand had not yet reached the target. Over the course of trials, participants adapted to the manipulation and pointed to a location between the visual target and the tactile buzz. We measured the perceived location of the hand by flashing a virtual pair of left and right hands before and after adaptation. Participants had to judge which hand they perceived closer to their body on the fronto-parallel plane. After adaptation, they judged the right hand, that corresponded to the hand used during adaptation, to be located further away from the body. We conclude that sensorimotor prediction of the consequences of hand movements shape sensory processing of hand location.
... Such Early studies of these neurons emphasized those with "congruent" discharge during execution 68 and observation conditions. Congruent neurons discharged during the same type of grasp 69 Rizzolatti et al., 1996), or retained the same preferred direction Emphasis on such congruent neurons led to the notion that they mediate understanding of 72 observed actions by mirroring their own activity during execution (di Pellegrino et al., 1992; 73 Rizzolatti and Craighero, 2004). 74 ...
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Many neurons in the premotor cortex show firing rate modulation whether the subject performs an action or observes another individual performing the same action. Although such "mirror neurons" have been thought to have highly congruent discharge during execution and observation, many if not most show non-congruent activity. Studies of such neuronal populations have shown that the most prevalent patterns of co-modulation—captured as neural trajectories—pass through subspaces which are shared in part, but in part are visited exclusively during either execution or observation. These studies focused on reaching movements for which the neural trajectories show comparatively simple dynamical motifs. But the neural dynamics of hand movements are more complex. We developed a novel approach to examine prevalent patterns of co-modulation during execution and observation of a task that involved reaching, grasping and manipulation. Rather than following neural trajectories in subspaces that contain their entire time course, we identified time series of instantaneous subspaces, sampled trajectory segments at the times of selected behavioral events, and projected each segment into the series of instantaneous subspaces. We found that instantaneous neural subspaces were partially shared between execution and observation in only one of three monkeys and were otherwise exclusive to one context or the other. Nevertheless, the patterns during execution and observation could be aligned with canonical correlation, indicating that though distinct, neural representations during execution and observation show dynamical similarity that may enable the nervous system to recognize particular actions whether performed by the subject or by another individual.
... However, it is only in the past 15 years that neurophysiological evidence started to accumulate . The most dramatic discovery was that 'mirror neurons' in the monkey ventral premotor cortex discharge during the execution of goal-directed hand movements and also when the monkey observes similar hand actions (Rizzolatti et al. 1996a). Another region in the monkey brain containing neurons specifically responsive to the sight of actions performed by others is in the STS (Perrett et al. 1989;Jellema et al. 2002). ...
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Humans, like other primates, are intensely social creatures. One of the main functions of our brains is to enable us to be as skilful in social interactions as we are in our interactions with the physical world. Any differences between human brains and those of our nearest relatives, the great apes, are likely to be linked to our unique achievements in social interaction and communication rather than our motor or perceptual skills. Unique to humans is the ability to mentalise (or mind read), that is to perceive and communicate mental states, such as beliefs and desires. A key problem facing neuroscience is to uncover the biological mechanisms underlying our ability to read other minds and to show how these mechanisms evolved. To solve this problem we need to do experiments in which people (or animals) interact with one another rather than behaving in isolation. Such experiments are now being conducted in increasing numbers and many of the leading exponents of such experiments have contributed to this volume. ‘The Neuroscience of Social Interactions’ will be an important step in uncovering the biological mechanisms underlying social interactions - undoubtedly one of the major programmes for neuroscience in the twenty-first century.
... There is neurophysiological evidence for a close relationship between the perception and performance of actions, at least on a fine temporal grain. Rizzolatti and his colleagues have shown that there are neurons in the premotor cortex of the monkey that fire both when a monkey performs a goal-directed action and when the monkey observes another monkey (or human experimenter) perform the same action (Rizzolatti, Fadiga, Gallese, & Fogassi, 1996). The actions represented by these cells can be quite specific. ...
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Events can be understood in terms of their temporal structure. The authors first draw on several bodies of research to construct an analysis of how people use event structure in perception, understanding, planning, and action. Philosophy provides a grounding for the basic units of events and actions. Perceptual psychology provides an analogy to object perception: Like objects, events belong to categories, and, like objects, events have parts. These relationships generate 2 hierarchical organizations for events: taxonomies and partonomies. Event partonomies have been studied by looking at how people segment activity as it happens. Structured representations of events can relate partonomy to goal relationships and causal structure; such representations have been shown to drive narrative comprehension, memory, and planning. Computational models provide insight into how mental representations might be organized and transformed. These different approaches to event structure converge on an explanation of how multiple sources of information interact in event perception and conception.
... Los estudios primarios están relacionados con la comprensión de la acción y con una integración desde las diferentes áreas cerebrales para generar un nuevo aprendizaje y una nueva conexión neuronal. 15,16 No obstante, las MN también han captado la atención y la imaginación de neurocientíficos, psicólogos y filósofos, ya que también han sido implicadas en simulación corporal, 17 empatía, 18 reconocimiento de emociones, 19 lectura de intenciones, 20 adquisición del lenguaje, 21 evolución del lenguaje, 22 comunicación manual, 16 procesamiento del lenguaje de señas, 23 percepción del habla, 24 producción del habla, 25 procesamiento de música, 26 orientación sexual 27 y experiencia estética. 28 Además, se ha sugerido que la disfunción de MN contribuye a una serie de trastornos, incluidos el autismo, la esquizofrenia, el síndrome de Down, la esclerosis múltiple, la adicción al cigarrillo y la obesidad. ...
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RESUMEN Introducción: se ha sugerido que las neuronas espejo (MN, por sus siglas en inglés) desempeñan un papel fundamental en los procesos sociales de orden superior, incluido el aprendizaje motor, la comprensión de la acción, el aprendi-zaje por imitación, la toma de perspectiva, la comprensión de las emociones faciales y la empatía. Sin embargo, aún no existen estudios que evalúen y demuestren un papel importante de las MN en el aprendizaje por imitación en instancias como la simulación clínica para estudiantes universitarios de carreras de ciencias de la salud. Objetivo: describir cómo algunos mecanismos neuronales pueden contribuir al aprendizaje basado en la imitación y discutir su papel en la simulación clínica. Material y métodos: se realizó una búsqueda sistemática en bases de datos electró-nicas, con el fin de recopilar la literatura disponible sobre el tema. Resultados: se incluyeron 22 artículos después de aplicar los criterios de inclusión y exclusión. Siete de ellos abordaron el proceso de aprendizaje a través de neuronas espejo, siete lo hicieron a través de la imitación y dos más trataron el aprendizaje sensoriomotor a través de MN. Conclusiones: aunque parece que el sistema de las MN tiene implicaciones que podrían subyacer al aprendizaje basado en simulación clínica, se necesitan más estudios primarios con alto rigor metodológico y experimentos para medir la actividad del sistema de las MN en el aprendizaje basado en simulación clínica y su papel para establecer conclusiones más concretas. ABSTRACT Introduction: mirror neurons (MN) have been suggested to play a fundamental role in higher order social processes, including motor learning, understanding action, imitation learning, taking perspective, understanding facial emotions, and empathy. However, there are still no studies that evaluate and demonstrate an important role of MN in imitation learning in instances such as clinical simulation for university students of health-sciences majors. Objective: to describe how some neural mechanisms can contribute to imitation based learning and discuss their role in clinical simulation. Material and methods: a systematic search was carried out in electronic databases, in order to compile the available literature on the subject. Results: twenty-two articles were included after applying the inclusion and exclusion criteria. Seven articles addressed the learning process through mirror neurons, seven addressed learning through imitation and two more treated sensorimotor learning through MN. Conclusions: although it seems that the MN system has implications that could underlie clinical simulation-based learning, more primary studies with high methodological rigor and experiments are needed to measure the activity of the MN system in clinical simulation-based learning and its role in order to establish more concrete conclusions.
... Los estudios primarios están relacionados con la comprensión de la acción y con una integración desde las diferentes áreas cerebrales para generar un nuevo aprendizaje y una nueva conexión neuronal. 15,16 No obstante, las MN también han captado la atención y la imaginación de neurocientíficos, psicólogos y filósofos, ya que también han sido implicadas en simulación corporal, 17 empatía, 18 reconocimiento de emociones, 19 lectura de intenciones, 20 adquisición del lenguaje, 21 evolución del lenguaje, 22 comunicación manual, 16 procesamiento del lenguaje de señas, 23 percepción del habla, 24 producción del habla, 25 procesamiento de música, 26 orientación sexual 27 y experiencia estética. 28 Además, se ha sugerido que la disfunción de MN contribuye a una serie de trastornos, incluidos el autismo, la esquizofrenia, el síndrome de Down, la esclerosis múltiple, la adicción al cigarrillo y la obesidad. ...
... She regarded prayer as a dynamic process of the spiritual ascent of a person who, having passed through seven stages, all of which correspond to a certain degree of prayer, reaches the centre or as she calls it "the palace". This incredibly beautiful palace shining with diamonds is our spiritual inner world (Ribot, 1896). ...
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Background. The analysis is based on the thesis that prayer is not only dialogue in the context of man-God relationship but also a kind of mental state characterized by empathy and synergy of both exclusively human and divine. The main prayer's nuance of meaning-appeal, response, insight, on the one hand, form the dialogue as an opportunity to accept the existence of God in the subjective dimension of I-You, and on the other-contribute to transcendence and self-construction of personality. In this prayer process, the perception of self as Other and Other as self is carried out. our study The purpose of the research is to find out Self-Other relationship in the context of prayer, its psychological and religious capabilities, which reveal its functional aspects Methods. Analysis, comparison, synthesis, generalization and systematization Results. Prayer, opening the prospects of overcoming existential problems, firstly, satisfies the spiritual needs of man through dialogue with the Deity. Secondly, it forms sensitivity to the Other. Thirdly, it performs the function of centering of consciousness, thanks to which centering individuality is transformed into an integral personality. Attention is paid to the functional relationship of faith and prayer, their role in constructing the prayer experience as a kind of empathic life strategy and as a religious and spiritual space of coexistence of I and Other. Prayer experience promotes forming the basis, life affirmation of a person and contains significant psychotherapeutic potential, as its carrier is aimed at effective and practical overcoming of the finiteness of human existence in the pursuit of eternal, absolute existence. Conclusion. Prayer as a dialogue motivates the activation of a people's essential forces, enriching their experience, personal growth and optimizing spiritual self-transcendence. Prayer intentionally contains the Other in itself, ensures co-involvement with Him. Its intensity depends on the emotional palette, which enhances the dialogue and constructs self. Prayer not only "focuses" consciousness, but also acts as a powerful means of centring it. This ability of prayer, enabling the process of unity of the human and the Divine, contributes to the formation of a harmonious, integral personality. The essence of prayer is most clearly revealed in prayer¬empathy when the boundaries of self are overcome and problems, the feelings of the Other find a response in self, a kind of personal overflow takes place. Prayer contains a significant psychotherapeutic potential, helps to overcome difficult life situations and contributes to the improvement of mental health, and during periods of life crises, ensure moral survival and affirmation of a personality.
... Transcranial magnetic stimulation (TMS) and motor-evoked potentials (MEPs) can be used to assess CE; these techniques have been used to report pain-induced reduction in CE [12][13][14][15]. The hypertonic saline-induced decrease in CE due to pain in an experimental skeletal muscle pain model was inhibited by action observation and motor imagery tasks, which exhibit a facilitating effect on CE [16,17]. However, action observation and motor imagery showed effects on the nervous system, but not on the pain itself. ...
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Skeletal muscle pain can reduce corticomotor excitability (CE). We examined the effects of mirror therapy (MT), which can increase CE, on skeletal muscle pain. Fifteen healthy adults were injected with hypertonic saline (5.8% NaCl, 0.2 mL) into the first dorsal interosseous muscle (FDI) of the right hand to induce an experimental skeletal muscle pain condition and then divided into pain + MT and pain groups. The pain + MT group observed their left index finger abducting and adducting for 4 min after injection, creating the illusion that they were performing the movement with their right index finger. However, the pain group remained at rest. CE was assessed by measuring motor-evoked potentials (MEPs) of the right FDI through transcranial magnetic stimulation. Pain was assessed using a numerical rating scale (NRS). MEP amplitudes were significantly higher in the pain + MT group; the same trend was observed after MT. NRS scores were significantly lower in the pain + MT group from 60 to 390 s after injection; the time to pain disappearance was also shorter. MT could decrease CE reduction and alleviate skeletal muscle pain. MT may be effective in preventing plastic changes in the neural system associated with skeletal muscle pain and providing early pain relief.
... La observación de una acción induce la activación del mismo circuito nervioso destinado a controlar su ejecución, o sea, la automática simulación de la misma acción en el cerebro del observador. Se ha sugerido que este mecanismo de simulación puede estar en la base de una forma implícita de comprender las acciones ajenas Rizzolatti et al., 1996;vedi anche Gallese, 2000vedi anche Gallese, , 2001vedi anche Gallese, , 2003avedi anche Gallese, , 2003bvedi anche Gallese, , 2005avedi anche Gallese, , 2005bvedi anche Gallese, , 2006Gallese, Keysers y Rizzolatti, 2004;Rizzolatti y Craighero, 2004). Cuando las neuronas espejo se activan, lo mismo durante la ejecución como durante la observación de las acciones ajenas, especifican directamente la finalidad de la acción, de hecho se ha demostrado que las neuronas del área premotora F5, que codifican las acciones de agarrar con una mano (neuronas espejo incluidas), se activan al conseguir cierta finalidad (como agarrar un objeto) independientemente de los movimientos requeridos para conseguirlo, incluso cuando, usando un utensilio, los movimientos se oponen a los normalmente usados (Escola et al., 2004;Umiltà et al., 2006). ...
... Brain, 122: 199-208. Gallese, V., Fadiga, L., Fogassi, L., ve Rizzolatti, G. (1996). Action recognition in the premotor cortex. ...
... La identificació amb el que s'ha observat és un fenomen ocasionat per les neurones mirall. Aquesta es produeix en dues modalitats, una identificació motora (Rizzolatti et al., 1996) i una altra d'emocional (Rizzolatti i Craighero, 2004). En conseqüència, gràcies a l'activitat d'aquestes neurones, a partir de l'observació d'una emoció o una acció, en el subjecte que observa s'activen circuits neuronals similars a aquells que s'activarien si aquest realitzés la mateixa acció o sentís la mateixa emoció. ...
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According to the mirror mechanism the discharge of F5 mirror neurons of a monkey observing another individual performing an action is a motor representation of the observed action that may serve to understand or learn from the action. This hypothesis, if strictly interpreted, requires mirror neurons to exhibit an action tuning that is shared between action observation and execution. Due to insufficient data it remains contentious if this requirement is met. To fill in the gaps, we conducted an experiment in which identical objects had to be manipulated in three different ways in order to serve distinct action goals. Using three methods, including cross-task classification, we found that at most time points F5 mirror neurons did not encode observed actions with the same code underlying action execution. However, in about 20% of neurons there were time periods with a shared code. These time periods formed a distinct cluster and cannot be considered a product of chance. Population classification yielded non-shared coding for observed actions in the whole population, which was at times optimal and consistently better than shared coding in differentially selected subpopulations. These results support the hypothesis of a representation of observed actions based on a strictly defined mirror mechanism only for small subsets of neurons and only under the assumption of time-resolved readout. Considering alternative concepts and recent findings, we propose that during observation mirror neurons represent the process of a goal pursuit from the observer’s viewpoint. Whether the observer’s goal pursuit, in which the other’s action goal becomes the observer’s action goal, or the other’s goal pursuit is represented remains to be clarified. In any case, it may allow the observer to use expectations associated with a goal pursuit to directly intervene in or learn from another’s action.
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According to the mirror mechanism the discharge of F5 mirror neurons of a monkey observing another individual performing an action is a motor representation of the observed action that may serve to understand or learn from the action. This hypothesis, if strictly interpreted, requires mirror neurons to exhibit an action tuning that is shared between action observation and execution. Due to insufficient data it remains contentious if this requirement is met. To fill in the gaps, we conducted an experiment in which identical objects had to be manipulated in three different ways in order to serve distinct action goals. Using three methods, including cross-task classification, we found that at most time points F5 mirror neurons did not encode observed actions with the same code underlying action execution. However, in about 20% of neurons there were time periods with a shared code. These time periods formed a distinct cluster and cannot be considered a product of chance. Population classification yielded non-shared coding for observed actions in the whole population, which was at times optimal and consistently better than shared coding in differentially selected subpopulations. These results support the hypothesis of a representation of observed actions based on a strictly defined mirror mechanism only for small subsets of neurons and only under the assumption of time-resolved readout. Considering alternative concepts and recent findings, we propose that during observation mirror neurons represent the process of a goal pursuit from the observer’s viewpoint. Whether the observer’s goal pursuit, in which the other’s action goal becomes the observer’s action goal, or the other’s goal pursuit is represented remains to be clarified. In any case, it may allow the observer to use expectations associated with a goal pursuit to directly intervene in or learn from another’s action.
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According to the mirror mechanism the discharge of F5 mirror neurons of a monkey observing another individual performing an action is a motor representation of the observed action that may serve to understand or learn from the action. This hypothesis, if strictly interpreted, requires mirror neurons to exhibit an action tuning that is shared between action observation and execution. Due to insufficient data it remains contentious if this requirement is met. To fill in the gaps, we conducted an experiment in which identical objects had to be manipulated in three different ways in order to serve distinct action goals. Using three methods, including cross-task classification, we found that at most time points F5 mirror neurons did not encode observed actions with the same code underlying action execution. However, in about 20% of neurons there were time periods with a shared code. These time periods formed a distinct cluster and cannot be considered a product of chance. Population classification yielded non-shared coding for observed actions in the whole population, which was at times optimal and consistently better than shared coding in differentially selected subpopulations. These results support the hypothesis of a representation of observed actions based on a strictly defined mirror mechanism only for small subsets of neurons and only under the assumption of time-resolved readout. Considering alternative concepts and recent findings, we propose that during observation mirror neurons represent the process of a goal pursuit from the observer’s viewpoint. Whether the observer’s goal pursuit, in which the other’s action goal becomes the observer’s action goal, or the other’s goal pursuit is represented remains to be clarified. In any case, it may allow the observer to use expectations associated with a goal pursuit to directly intervene in or learn from another’s action.
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SUMMARY AND CONCLUSIONS 1. We recorded from single neurons in the dorsal bank and fundus of the anterior por- tion of the superior temporal sulcus, an area we term the superior temporal polysensory area (STP). Five macaques were studied under anesthesia ( N20) and immobilization in repeated recording sessions. 2. Almost all of the neurons were visually responsive, and over half responded to more than one sensory modality; 21% responded to visual and auditory stimuli, 17% re- sponded to visual and somesthetic stimuli, 17% were trimodal, and 41% were exclu- sively visual. 3. Almost all the visual receptive fields extended into both visual half-fields, and the majority approached the size of the visual field of the monkey, including both monoc- ular crescents. Somesthetic receptive fields were also bilateral and usually included most of the body surface. 4. Virtually all neurons responded better to moving visual stimuli than to stationary visual stimuli, and almost half were sensitive to the direction of movement. Several classes of directional neurons were found, including a) neurons selective for a single direction of movement throughout their receptive field, b) neurons selective for directions of move- ment radially symmetric about the center of gaze, and c) neurons selective for movement in depth. 5. The majority of neurons (70%) had lit- tle or no preference for stimulus size, shape, orientation, or contrast. The minority (30%) responded best to particular stimuli. Some of these appeared to be selective for faces. 6. The properties of most STP neurons, such as large receptive fields, sensitivity to movement, insensitivity to form, and poly- modal responsiveness, suggest that STP is more involved in orientation and spatial functions than in pattern recognition.
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Primates represent the most highly evolved order in the zoological system. Nevertheless, they have a number of communicatory signals that, similar to lower organisms, are not shaped by imitation from conspecifics but are genetically determined. Such signals include most nonverbal vocalizations (monkey calls; human laughing, crying, moaning), many facial expressions (monkey threatening face and submission grin, human smiling and frowning) as well as more complex display patterns, such as the genital display of the squirrel monkey, lip smacking of macaques, or chest beating of the gorilla. The motor control of innate and learned patterns relies on partially different neural mechanisms in the sense that there are a number of brain structures involved in the production of human speech, whistling, or gestural signing that are dispensable for the production of pain cries, smiling, or fist clenching during rage. These brain structures are superimposed on, not independent of, those controlling innate behavior.
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The concept is presented that human speech normally depends on two systems working harmoniously together. The one system is phylogenetically older, is subordinate, and is supplementary to the other. Anatomical and clinical considerations suggest that the older system normally functions through the newer by means of forebrain connections but that vestigial limbic speech is still possible through its brain stem connections. In rational and logical discourse, the neocortical system is predominant, but in times of emotional stress the limbic system may reclaim its old primacy. Lesions affecting speech provide an insight into the modus operandi of the two systems. The resulting model supplies an explanation for the discrepancies noted above in the neocortical theory of speech. Namely, lesions of limbic structures neighboring the neocortical speech center account for the lack of one to one mapping between neocortical lesions and the type and degree of aphasia; vestigial limbic speech appears as the selective preservation of emotionally charged or vulgar words and phrases; withdrawal of limbic facilitation produces mutism; and heightened limbic facilitation results in transient qualitative and quantitative improvement in the nonfluent dysphasias and may increase the rate and degree of ultimate recovery.
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If the present interpretations are valid, it is apparent that the speech of man has evolved not from the vocal responses of lower primates, but rather speech has developed de novo in man during his evolutionary development beyond the level of monkeys or, indeed, apparently, of the apes. From a neurologic view, the evolution of speech must represent the evolution of those mechanisms of the cerebrum located posteriorly in zones of cortex that function to analyze the information of the senses, to establish memories thereof, and to organize voluntary responses which proceed from these analyses or memories. The separate and distinct mechanisms of the cerebrum that control emotional and instinctive behavior still remain in the human to link us phylogenetically with the lower primate forebears. For the proper study of language, however, one would do well to focus on the physiologic properties of the posterior cerebrum of man himself and to avoid speculation as to similarities between the vocal responses of animals and speech in man.
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The mesial agranular frontal cortex that lies rostral to area 4 (F1) is formed by two distinct cytoarchitectonic areas: F3, located caudally, and F6, located rostrally. In the present experiments we investigated the organization of F3 and F6 by observing the motor responses evoked by their intracortical electrical microstimulation. Our main purpose was to find out whether the cytoarchitectonic subdivision of the mesial agranular frontal cortex into two areas has a physiological counterpart. The result showed that F3 (the caudal area) contains a complete motor representation with hindlimb movements located caudally, forelimb movements located centrally, and orofacial movements located rostrally. The great majority of limb movements involved proximal joints. With respect to F1, F3 showed the following functional characteristics: (1) lack of segregation between proximal and distal movements, (2) larger percentage of complex movements, and (3) higher excitability threshold. Movements were more difficult to elicit from F6 (the rostral area) than from F3. However, by using a longer stimulus train duration (100 ms) 39.3% of tested sites produced body movements. This percentage increased (50.5%) when the electrical stimulation was applied during monkey natural movements instead of when the monkey was still in its chair. Most of the evoked movements concerned the forelimb. More rarely, neck and upper face movements were observed. Unlike F1 and F3 where most movements were fast, slow movements were frequently observed with stimulation of F6. Many of them mimicked natural movements of the animal. Eye movements were evoked from F7 (superior area 6) but not from F6. An additional motor representation was found in the dorsocaudal part of area 24 (24d). This area is topographically organized with a forelimb representation located caudally and ventrally and a hindlimb representation located rostrally and dorsally. The excitability threshold of area 24d is higher than that of F1 and F3. Evoked movements were occasionally observed also after stimulation of area 24c. In conclusion, on the mesial cortical wall rostral to F1, there are at least three independent motor representations. On the basis of somatotopic organization and excitability properties, we propose that the term supplementary motor area (SMA-proper) should be reserved to F3.
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The agranular frontal cortex is formed by several distinct functional areas. There is no agreement, however, on its cytoarchitectonic organization. The aim of this study was to redefine the cytoarchitectonic organization of superior and mesial area 6 and the adjacent cingulate cortex in the macaque monkey. A particular goal was to find out whether the so-called supplementary motor area (SMA) is cytoarchitectonically different from the rest of area 6 and whether it can be considered as a single, independent cytoarchitectonic area. The results showed that, rostral to F1 (area 4), four architectonic areas can be recognized in the superior (dorsal) and mesial area 6. Two of them are located on mesial cortical surface (F3 caudally and F6 rostrally) and two on superior cortical convexity (F2 caudally and F7 rostrally). The main cytoarchitectonic features of the five identified areas can be summarized as follows. F1: (1) giant pyramidal cells organized in multiple rows. (2) columnar pattern extending from the white matter to the superficial layers, (3) low cellular density in the lower part of layer III. F3: (1) high cellular density in the lower part of layer III, which fuses with a dense Va, (2) columnar pattern present only in the deepest layer, (3) occasional presence of giant pyramidal cells in layer Vb. F6: (1) prominent layer V, (2) absence of sublayer Vb, (3) homogeneous cell density in superficial layers. F2: (1) thin row of medium-size pyramids in the lowest part of layer III, (2) columnar pattern extending to the superficial layers, (3) dense layer Va, (4) few, scattered giant pyramids in layer Vb. F7: (1) prominent layer V, (2) bipartite layer VI. Areas F1, F2, and F3, as defined cytoarchitectonically, coincided with the homonymous histochemical areas. The present data showed also that area 24 is formed by four subareas: 24a, b, c and d. Areas 24a ana b occupy the ventral part of area 24, whereas its dorsal part is formed by area 24c, located rostrally, and area 24d, located caudally. The following features distinguish area 24d from area 24c: (1) larger pyramidal cells in layer V, (2) presence of medium-size pyramidal cells in the lower part of layer III, (3) more prominent columnar pattern, (4) higher myelinization with the presence of an evident horizontal plexus. Mesial area 6 is usually considered as a single functional entity (SMA). Our findings show that this cortical region is formed by two distinct cytoarchitectonic areas. In the following article (Luppino et al. 1991: J. Comp. Neurol 311:463–482) physiological evidence is presented that the SMA, as classically defined, corresponds to F3, whereas F6 is an independent functional area.
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Neuronal tracers (diamidino yellow or wheat germ agglutinin conjugated with horseradish peroxidase) were injected in the arm representations of area 6a alpha (mesial surface, area F3), in the arm representation of area 6a beta (mesial surface) as well as in the eye field of area 6a beta (dorso-medial surface). The results showed that the arm representation of area F3 receives topographically organized afferents from motor and premotor areas (areas F1, F2, F4 and F5). A further connection was found with that part of cingulate cortex that sends projections to the spinal cord. In contrast, the arm representation of area 6a beta receives afferents chiefly from area F5, the prefrontal cortex and that part of cingulate sulcus which has few, if any, connections with the spinal cord. No connections were found with the precentral motor cortex (area F1). The area 6a beta eye field receives afferents mostly from the frontal eye field. Further connections are with the prefrontal cortex and cingulate gyrus. It is suggested that the so called "low level" motor functions of supplementary motor area are due to the activity of area F3, whereas the so called "high level" motor functions depend upon an independent area located in area 6a beta.
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Cognition and comportment are subserved by interconnected neural networks that allow high-level computational architectures including parallel distributed processing. Cognitive problems are not resolved by a sequential and hierarchical progression toward predetermined goals but instead by a simultaneous and interactive consideration of multiple possibilities and constraints until a satisfactory fit is achieved. The resultant texture of mental activity is characterized by almost infinite richness and flexibility. According to this model, complex behavior is mapped at the level of multifocal neural systems rather than specific anatomical sites, giving rise to brain-behavior relationships that are both localized and distributed. Each network contains anatomically addressed channels for transferring information content and chemically addressed pathways for modulating behavioral tone. This approach provides a blueprint for reexploring the neurological foundations of attention, language, memory, and frontal lobe function.
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Single neurons were recorded from the rostral part of the agranular frontal cortex (area 6a beta) in awake, partially restrained macaque monkeys. In the medialmost and mesial sectors of this area, rostral to the supplementary motor area, neurons were found which were activated during arm reaching-grasping movements. These neurons ("reaching-grasping neurons") did not appear to be influenced by how the objects were grasped nor, with some exceptions, by where they were located. Their activity changed largely prior to the arm movement and continued until the end of it. The premovement modulation (excitatory or inhibitory) could start with stimulus presentation, with the saccade triggered by the stimulus or after stimulus fixation. The distance of the stimulus from the monkey was an important variable for activating many neurons. About half of the recorded neurons showed a modulation of the same sign during movement and premovement period. The other half showed an increase/decrease in activity which was of the opposite sign during movement and premovement period or part of it. In this last case the discharge changes were of the same sign when the stimulus was close to the monkey and when the monkey moved its arm to reach the objects, whereas they were of opposite sign when the stimulus was outside the animal's reach. Microstimulation of area 6a beta and the reconstruction of the locations of eye movement and arm movement related cells showed that the arm field was located more medially (and mesially) than the eye field described by Schlag and Schlag-Rey (1987). It is suggested that, unlike inferior area 6, which is mostly involved in selection of effectors on the basis of the physical properties of the objects and their spatial location (Rizzolatti and Gentilucci 1988), area 6a beta plays a role in the preparation of reaching-grasping arm movements and in their release when the appropriate conditions are set.
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The presence of neurons in macaque temporal cortex and amygdala which fire selectively in response to social stimuli has been demonstrated by several investigators. The extent to which such neuronal populations may respond to a broad range of social features, including expressive movements and interactions, has not been fully explored due to the difficulty of presenting such complex stimuli in a controlled fashion. We describe a method for presenting moving segments of macaque behavior, visual and auditory, to animal subjects during single unit recording. The method permits a broad range of stimuli to be used both as probes and as controls. In addition, a novel technique for monitoring eye position in alert macaque subjects is described. We present results from the medial amygdala and adjacent cortex, demonstrating that neurons in these regions respond selectively to features of the social environment.
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We have examined the circuitry connecting the posterior parietal cortex with the frontal lobe of rhesus monkeys. HRP-WGA and tritiated amino acids were injected into subdivisions 7m, 7a, 7b, and 7ip of the posterior parietal cortex, and anterograde and retrograde label was recorded within the frontal motor and association cortices. Our main finding is that each subdivision of parietal cortex is connected with a unique set of frontal areas. Thus, area 7m, on the medial parietal surface, is interconnected with the dorsal premotor cortex and the supplementary motor area, including the supplementary eye field. Within the prefrontal cortex, area 7m's connections are with the rostral sector of the frontal eye field (FEF), the dorsal bank of the principal sulcus, and the anterior bank of the inferior arcuate sulcus (Walker's area 45). In contrast, area 7a, on the posterior parietal convexity, is not linked with premotor regions but is heavily interconnected with the rostral FEF in the anterior bank of the superior arcuate sulcus, the dorsolateral prefrontal convexity, the rostral orbitofrontal cortex, area 45, and the fundus and adjacent cortex of the dorsal and ventral banks of the principal sulcus. Area 7b, in the anterior part of the posterior parietal lobule, is interconnected with still a different set of frontal areas, which include the ventral premotor cortex and supplementary motor area, area 45, and the external part of the ventral bank of the principal sulcus. The prominent connections of area 7ip, in the posterior bank of the intraparietal sulcus, are with the supplementary eye field and restricted portions of the ventral premotor cortex, with a wide area of the FEF that includes both its rostral and caudal sectors, and with area 45. All frontoparietal connections are reciprocal, and although they are most prominent within a hemisphere, notable interhemispheric connections are also present.
Article
The rostral part of the agranular frontal cortex (area 6) can be subdivided on the basis of its cytoarchitecture, enzymatic properties, and connections into two large sectors: a superior region, lying medial to the spur of the arcuate sulcus, and an inferior region, lying lateral to it. In this study we traced the afferent and efferent connections of the inferior region of area 6 by injecting small amounts of wheat germ agglutinin conjugated to horseradish peroxidase (WGA‐HRP) and fluorescent tracers (fast blue and diamidino yellow) into restricted parts of inferior area 6 and in physiologically determined fields of area 4. There is an ordered topographic pattern of connections between inferior area 6 and area 4. The region near the spur of the arcuate sulcus (hand field) projects to the area 4 hand field while the lateral part of inferior area 6 (mouth field) is connected with the corresponding field in area 4. The organization of the connections between the two fields is, however, different. The hand fields in area 6 and 4 have direct reciprocal projections, whereas the mouth field in the postarcuate cortex relays information to area 4 via a zone intermediate between the arcuate and the central sulcus. This zone corresponds to the cytochrome oxidase area F4 (Matelli, Luppino, and Rizzolatti: Behav . Brain Res . 18 : 125–137, '85). The inferior area 6 also has topographically organized connections with the supplementary motor area. The inferior area 6 receives and sends fibers to a series of discrete cortical areas located in the lower cortical moiety (Sanides: The Structure and Function of the Nervous Tissue , Vol. 5. New York: Academic Press, pp 329–453, '72). These areas that form a broad ring around the central sulcus are the ventral bank of the principal sulcus and the adjacent area 46, the precentral operculum (PrOC), area SII (Jones and Burton: J. Comp. Neurol. 168 :197–248, '76), the parietal operculum, and the rostral part of the inferior parietal lobule including the lower bank of the intraparietal suclus. Finall, the inferior area 6 has sparse but consistent connections with insular and cingulate cortices. The functional significance of this complex pattern of connections is discussed.
Article
The functional properties of neurons located in the rostral part of inferior area 6 were studied in awake, partially restrained macaque monkeys. The most interesting property of these neurons was that their firing correlated with specific goal-related motor acts rather than with single movements made by the animal. Using the motor acts as the classification criterion we subdivided the neurons into six classes, four related to distal motor acts and two related to proximal motor acts. The distal classes are: "Grasping-with-the-hand-and-the-mouth neurons", "Grasping-with-the-hand neurons", "Holding neurons" and "Tearing neurons". The proximal classes are: "Reaching neurons" and "Bringing-to-the-mouth-or-to-the-body neurons". The vast majority of the cells belonged to the distal classes. A particularly interesting aspect of distal class neurons was that the discharge of many of them depended on the way in which the hand was shaped during the motor act. Three main groups of neurons were distinguished: "Precision grip neurons", "Finger prehension neurons", "Whole hand prehension neurons". Almost the totality of neurons fired during motor acts performed with either hand. About 50% of the recorded neurons responded to somatosensory stimuli and about 20% to visual stimuli. Visual neurons were more difficult to trigger than the corresponding neurons located in the caudal part of inferior area 6 (area F4). They required motivationally meaningful stimuli and for some of them the size of the stimulus was also critical. In the case of distal neurons there was a relationship between the type of prehension coded by the cells and the size of the stimulus effective in triggering the neurons. It is proposed that the different classes of neurons form a vocabulary of motor acts and that this vocabulary can be assessed by somatosensory and visual stimuli.
Article
Two series of experiments are reported in this paper. The first concerns the movement representation in the macaque inferior area 6, the second the functional properties of neurons located in the caudal part of this area (histochemical area F4). By combining single neuron recording and intracortical microstimulation, we found that inferior area 6 is somatotopically organized. The axio-proximal movements are represented caudally, the distal movements are represented near the arcuate sulcus. The mouth field is located laterally, the hand field medially. There is no leg field. A comparison between neuron properties and histochemical characteristics of inferior area 6 showed that the proximal movements representation includes most of area F4, whereas the distal movements representation corresponds to area F5 and to the rostral part of F4. Neurons located in that part of F4 where proximal movements are represented respond very well to tactile stimuli. They have large receptive fields mostly located on the face and on the upper part of the body. A large number of these neurons respond to visual stimuli. Objects approaching the animal are particularly effective. The tactile and the visual receptive fields are in register. The most represented movements are reaching movements, movements bringing the hand to the mouth or to the body and facial movements. There is a congruence between location of visual fields and preferred arm movements. It is argued that the receptive field arrangement and the response properties are more complex in area F4 than in the primary motor cortex and that area F4 neurons are involved in the control of arm movements towards different space sectors.
Article
The concept of a separate premotor cortical field involved in the cerebral control of movement went out of favor among neurophysiologists during the quarter century from 1952 to 1977, but recent studies have led to its rehabilitation. The premotor cortex appears to be one of at least three fields within the motor cortex, two others being the primary motor cortex and the supplementary motor cortex. Several proposals have been presented concerning the functional specializations of the premotor cortex. Although no specific hypotheses have very strong support at present, the best evidence favors a role for premotor cortex in the preparation for and the sensory guidance of movement.
Article
To investigate the functions of the amygdala in visual information processing and in emotional and social responses, recordings were made from single neurons in the amygdala of the monkey. A population of neurons (40 of more than 1000 recorded in 4 monkeys) was investigated which responded primarily to faces. These neurons typically (1) responded to some human or monkey faces, which were presented to the monkey through a large aperture shutter so that response latencies could be measured, or were simply shown to the monkey, (2) responded to 2-dimensional representations of these faces, as well as to real 3-dimensional faces, (3) had no responses or only small (less than half maximum) responses to gratings, simple geometrical, other complex 3-D stimuli, or to arousing and aversive stimuli, (4) had response latencies of 110-200 ms, (5) were located in the basal accessory nucleus of the amygdala, (6) responded differently to different faces, as shown by measures of d', and could thus over a population of such neurons code information useful for making different responses to different individuals, (7) could in some cases (9/11 tested) respond to parts of faces, and (8) in a few cases (4/19 tested) responded more to a face which produced an emotional response. A comparison made in three monkeys of the responses of these neurons with the responses of 77 neurons with face-selective responses recorded in the cortex of the superior temporal sulcus (STS) showed that the amygdaloid neurons had longer response latencies (110-200 compared to 90-140 ms), and were in some respects more selective in their responses to different faces. It is suggested that the deficits in social and emotional behavior produced by amygdala lesions could be due in part to damage to a neuronal system specialized in utilizing information from faces so that appropriate social and emotional responses can be made to different individuals.
Article
The projections to the frontal cortex from the various subdivisions of the posterior parietal region in the rhesus monkey were studied by means of autoradiographic technique. The rostral superior parietal lobule (area PE) projects to the dorsal areas 4 and 6 on the lateral surface of the frontal lobe as well as to the supplementary motor area (MII) on its medial surface. The caudal area PE sends its connections to dorsal area 6 and MII. The projections from the medial parietal cortex (areas PEc and PGm) are similar to those of the superior parietal lobule but they tend to concentrate in the more rostral part of dorsal area 6, MII, and in the cingulate gyrus (area 24). The most caudal part of the medial parietal cortex also projects to area 8. The anteriormost part of the inferior parietal lobule (area PF) projects to the ventral area 6, including the caudal bank of the lower branch of the arcuate sulcus, to the ventral area 46 below the sulcus principalis, and to the frontal and pericentral opercular cortex. The middle inferior parietal lobule (areas PFG and PG) projects to the ventral part of area 46 and area 8, whilst the posteriormost inferior parietal lobule (caudal PG and area Opt) is connected with both dorsal and ventral area 46, dorsal area 8, as well as the anteriormost dorsal area 6, and the cingulate gyrus (area 24).
Article
The frontal agranular cortex consists of at least three major subdivisions: the precentral motor cortex, the supplementary motor cortex, and the premotor cortex. Of these, the premotor cortex is by far the least intensively studied by neurophysiological methods in spite of recent evidence that it is important in higher order aspects of the cerebral control of movement ( Moll and Kuypers , 1977; Roland et al., 1980b ; Halsband and Passingham , 1982). We have accordingly studied neuronal activity in the premotor cortex of Rhesus macaques operantly conditioned to perform a visually guided motor task. We have concluded that many premotor cortex neurons appear to reflect motor set. Further, these and other premotor cortex neurons show activity patterns during and before the execution of an abstractly guided movement that are strikingly similar to what has been observed in association with movements made directly to visuospatial targets. Finally, a functional relationship of some premotor cortex unit activity to the execution of voluntary movement is supported by its close temporal correlation with the onset of movement, a significant correlation with the acceleration of the limb in some cases, and a uniform specificity for movement execution rather than the visuospatial cues that guide the movement.
Article
Single neurons were recorded from the "mouth area" of the postarcuate cortex in macaque monkeys. According to their responses to somatosensory stimuli and their activity during the animal's movements, the neurons were subdivided into three classes: (1) neurons activated only by external stimuli; (2) neurons showing an enhancement of their response when a specific motor act followed the external stimulation; (3) neurons responding reliably to external stimuli only if the stimulation triggered a specific, related motor act. Of the recorded neurons, 50% also responded to visual stimuli.
Article
Grasping requires coding of the object's intrinsic properties (size and shape), and the transformation of these properties into a pattern of distal (finger and wrist) movements. Computational models address this behavior through the interaction of perceptual and motor schemas. In monkeys, the transformation of an object's intrinsic properties into specific grips takes place in a circuit that is formed by the inferior parietal lobule and the inferior premotor area (area F5). Neurons in both these areas code size, shape and orientation of objects, and specific types of grip that are necessary to grasp them. Grasping movements are coded more globally in the inferior parietal lobule, whereas they are more segmented in area F5. In humans, neuropsychological studies of patients with lesions to the parietal lobule confirm that primitive shape characteristics of an object for grasping are analyzed in the parietal lobe, and also demonstrate that this 'pragmatic' analysis of objects is separated from the 'semantic' analysis performed in the temporal lobe.