The Monkey Mirror

Povinelli believes that rhesus monkeys, in contrast, are simply engaging in normal behaviors like grooming and attempting to socialize with the mirror monkeys. He also maintains that the monkeys have not passed the mark test because they still have not recognized a traditional mark of dye or a sticker. Monkeys find the implants to be more interesting than traditional marks, he believes. Populin hopes to conduct further studies with the rhesus monkeys, using them to examine human conditions like gaze aversion, in which one person avoids eye contact with another, even during a conversation.

The argument over whether monkeys really recognize their mirror images is not just a narrow debate among primatologists. It also has broad implications for evolutionary theory. The clash is over how self-awareness evolved: The ability of monkeys to recognize themselves in a mirror might indicate a level of self-awareness that supports the idea of a gradual development of that ability along the evolutionary tree.

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Reports on mirror neurons have been widely published [24] and confirmed [25] with mirror neurons found in both inferior frontal and inferior parietal regions of the brain.

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Recently, evidence from functional neuroimaging strongly suggests that humans have similar mirror neurons systems: Not surprisingly, these brain regions include those found in the macaque monkey [1] However, functional magnetic resonance imaging fMRI can examine the entire brain at once and suggests that a much wider network of brain areas shows mirror properties in humans than previously thought.

These additional areas include the somatosensory cortex and are thought to make the observer feel what it feels like to move in the observed way. The most common theory behind the origin of mirror neuron is the genetic account which suggests that the mirrorness of mirror neurons is due primarily to heritable genetic factors and that the genetic predisposition to develop Mirror neuron evolved because they facilitate action understanding.

The other theories as to the origin of mirror neurons include Associative Learning , Canalization and Exaptation.

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The first animal in which researchers have studied mirror neurons individually is the macaque monkey. In these monkeys, mirror neurons are found in the inferior frontal gyrus region F5 and the inferior parietal lobule. Mirror neurons are believed to mediate the understanding of other animals' behaviour. For example, a mirror neuron which fires when the monkey rips a piece of paper would also fire when the monkey sees a person rip paper, or hears paper ripping without visual cues.

These properties have led researchers to believe that mirror neurons encode abstract concepts of actions like 'ripping paper', whether the action is performed by the monkey or another animal. The function of mirror neurons in macaques remains unknown. Adult macaques do not seem to learn by imitation. Recent experiments by Ferrari and colleagues suggest that infant macaques can imitate a human's face movements, though only as neonates and during a limited temporal window. In adult monkeys, mirror neurons may enable the monkey to understand what another monkey is doing, or to recognize the other monkey's action.

It is not normally possible to study single neurons in the human brain, so most evidence for mirror neurons in humans is indirect. Brain imaging experiments using functional magnetic resonance imaging fMRI have shown that the human inferior frontal cortex and superior parietal lobe are active when the person performs an action and also when the person sees another individual performing an action. It has been suggested that these brain regions contain mirror neurons, and they have been defined as the human mirror neuron system. Neuropsychological studies looking at lesion areas that cause action knowledge, pantomime interpretation, and biological motion perception deficits have pointed to a causal link between the integrity of the inferior frontal gyrus and these behaviours.

A study published in April reports recordings from single neurons with mirror properties in the human brain. The patients had been implanted with intracranial depth electrodes to identify seizure foci for potential surgical treatment. Electrode location was based solely on clinical criteria; the researchers, with the patients' consent, used the same electrodes to "piggyback" their research.

The Monkey in the Mirror

The researchers found a small number of neurons that fired or showed their greatest activity both when the individual performed a task and when they observed a task. Other neurons had anti-mirror properties, that is, they responded when the participant performed an action but were inhibited when the participant saw that action. The mirror neurons found were located in the supplementary motor area and medial temporal cortex other brain regions were not sampled.

For purely practical reasons, these regions are not the same as those in which mirror neurons had been recorded from in the monkey: On the other hand, no one has to date looked for mirror neurons in the supplementary motor area or the medial temporal lobe in the monkey. Together, this therefore does not suggest that humans and monkeys have mirror neurons in different locations, but rather that they may have mirror neurons both in the ventral premotor cortex and inferior parietal lobe, where they have been recorded in the monkey, and in the supplementary motor areas and medial temporal lobe, where they have been recorded from in human — especially because detailed human fMRI analyses suggest activity compatible with the presence of mirror neurons in all these regions.

Another study has suggested that human beings don't necessarily have more mirror neurons than monkeys, but instead that there is a core set of mirror neurons used in action observation and execution. However, for other proposed functions of mirror neurons the mirror system may have the ability to recruit other areas of the brain when doing its auditory, somatosensory, and affective components. Although many in the scientific community have expressed excitement about the discovery of mirror neurons, there are scientists who have expressed doubts about both the existence and role of mirror neurons in humans.

According to scientists such as Hickok, Pascolo, and Dinstein, it is not clear whether mirror neurons really form a distinct class of cells as opposed to an occasional phenomenon seen in cells that have other functions , [40] and whether mirror activity is a distinct type of response or simply an artifact of an overall facilitation of the motor system.

In , Ilan Dinstein et al. In , Lingnau et al. They concluded that there was a significant asymmetry between the two processes that indicated that mirror neurons do not exist in humans. They stated "Crucially, we found no signs of adaptation for motor acts that were first executed and then observed. Failure to find cross-modal adaptation for executed and observed motor acts is not compatible with the core assumption of mirror neuron theory, which holds that action recognition and understanding are based on motor simulation.

In , Greg Hickok published an extensive argument against the claim that mirror neurons are involved in action-understanding: However, despite its widespread acceptance, the proposal has never been adequately tested in monkeys, and in humans there is strong empirical evidence, in the form of physiological and neuropsychological double- dissociations, against the claim. Vladimir Kosonogov sees another contradiction.

The proponents of mirror neuron theory of action understanding postulate that the mirror neurons code the goals of others actions because they are activated if the observed action is goal-directed. However, the mirror neurons are activated only when the observed action is goal-directed object-directed action or a communicative gesture, which certainly has a goal too. How do they "know" that the definite action is goal-directed? At what stage of their activation do they detect a goal of the movement or its absence?

In his opinion, the mirror neuron system can be activated only after the goal of the observed action is attributed by some other brain structures. Neurophilosophers such as Patricia Churchland have expressed both scientific and philosophical objections to the theory that mirror neurons are responsible for understanding the intentions of others. In chapter 5 of her book, Braintrust, Churchland points out that the claim that mirror neurons are involved in understanding intentions through simulating observed actions is based on assumptions that are clouded by unresolved philosophical issues.

She makes the argument that intentions are understood coded at a more complex level of neural activity than that of individual neurons. Churchland states that "A neuron, though computationally complex, is just a neuron. It is not an intelligent homunculus. If a neural network represents something complex, such as an intention [to insult], it must have the right input and be in the right place in the neural circuitry to do that". Recently, Cecilia Heyes Professor of Experimental Psychology, Oxford has advanced the theory that mirror neurons are the byproduct of associative learning as opposed to evolutionary adaptation.

She argues that mirror neurons in humans are the product of social interaction and not an evolutionary adaptation for action-understanding. In particular, Heyes rejects the theory advanced by V. Ramachandran that mirror neurons have been "the driving force behind the great leap forward in human evolution. Two closely related models postulate that mirror neurons are trained through Hebbian [47] or Associative learning [48] [49] [50] see Associative Sequence Learning. However, if premotor neurons need to be trained by action in order to acquire mirror properties, it is unclear how newborn babies are able to mimic the facial gestures of another person imitation of unseen actions , as suggested by the work of Meltzoff and Moore.

One possibility is that the sight of tongue protrusion recruits an innate releasing mechanism in neonates. Careful analysis suggests that 'imitation' of this single gesture may account for almost all reports of facial mimicry by new-born infants. Many studies link mirror neurons to understanding goals and intentions. The IPL has long been recognized as an association cortex that integrates sensory information. The monkeys watched an experimenter either grasp an apple and bring it to his mouth or grasp an object and place it in a cup.

Only the type of action, and not the kinematic force with which models manipulated objects, determined neuron activity. It was also significant that neurons fired before the monkey observed the human model starting the second motor act bringing the object to the mouth or placing it in a cup. Therefore, IPL neurons "code the same act grasping in a different way according to the final goal of the action in which the act is embedded". Another possible function of mirror neurons would be facilitation of learning.

The mirror neurons code the concrete representation of the action, i. This would allow us to simulate to repeat internally the observed action implicitly in the brain to collect our own motor programs of observed actions and to get ready to reproduce the actions later. It is implicit training. This happens due to associative learning processes.

The more frequently a synaptic connection is activated, the stronger it becomes. Stephanie Preston and Frans de Waal , [53] Jean Decety , [54] [55] and Vittorio Gallese [56] [57] and Christian Keysers [3] have independently argued that the mirror neuron system is involved in empathy. A large number of experiments using fMRI, electroencephalography EEG and magnetoencephalography MEG have shown that certain brain regions in particular the anterior insula , anterior cingulate cortex , and inferior frontal cortex are active when people experience an emotion disgust, happiness, pain, etc.

More recently, Christian Keysers at the Social Brain Lab and colleagues have shown that people who are more empathic according to self-report questionnaires have stronger activations both in the mirror system for hand actions [66] and the mirror system for emotions, [63] providing more direct support for the idea that the mirror system is linked to empathy.

Some researchers observed that the human mirror system does not passively respond to the observation of actions but is influenced by the mindset of the observer. Ramachandran has speculated that mirror neurons may provide the neurological basis of human self-awareness. I also speculated that these neurons can not only help simulate other people's behavior but can be turned 'inward'—as it were—to create second-order representations or meta-representations of your own earlier brain processes.

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This could be the neural basis of introspection, and of the reciprocity of self awareness and other awareness. There is obviously a chicken-or-egg question here as to which evolved first, but The main point is that the two co-evolved, mutually enriching each other to create the mature representation of self that characterizes modern humans". In humans, functional MRI studies have reported finding areas homologous to the monkey mirror neuron system in the inferior frontal cortex, close to Broca's area, one of the hypothesized language regions of the brain.

Mirror neurons have been said to have the potential to provide a mechanism for action-understanding, imitation-learning, and the simulation of other people's behaviour. Such speech repetition occurs automatically, fast [73] and separately in the brain to speech perception. Further evidence for this link comes from a recent study in which the brain activity of two participants was measured using fMRI while they were gesturing words to each other using hand gestures with a game of charades — a modality that some have suggested might represent the evolutionary precursor of human language.

Analysis of the data using Granger Causality revealed that the mirror-neuron system of the observer indeed reflects the pattern of activity in the motor system of the sender, supporting the idea that the motor concept associated with the words is indeed transmitted from one brain to another using the mirror system [78].

The mirror neuron system seems to be inherently inadequate to play any role in syntax , given that this definitory property of human languages which is implemented in hierarchical recursive structure is flattened into linear sequences of phonemes making the recursive structure not accessible to sensory detection [79].

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The term is commonly used to refer to cases in which an individual, having observed a body movement, unintentionally performs a similar body movement or alters the way that a body movement is performed. Automatic imitation rarely involves overt execution of matching responses.

Instead the effects typically consist of reaction time, rather than accuracy, differences between compatible and incompatible trials. Research reveals that the existence of automatic imitation, which is a covert form of imitation, is distinct from spatial compatibility. It also indicates that, although automatic imitation is subject to input modulation by attentional processes, and output modulation by inhibitory processes, it is mediated by learned, long-term sensorimotor associations that cannot be altered directly by intentional processes.

Many researchers believe that automatic imitation is mediated by the mirror neuron system. For example, if the task is to maintain posture, people do it worse when they listen to sentences like this: This phenomenon may be due to the fact that during action perception there is similar motor cortex activation as if a human being performed the same action mirror neurons system. The integration of research on motor mimicry and automatic imitation could reveal plausible indications that these phenomena depend on the same psychological and neural processes.

Preliminary evidence however comes from studies showing that social priming has similar effects on motor mimicry. Nevertheless, the similarities between automatic imitation, mirror effects, and motor mimicry have led some researchers to propose that automatic imitation is mediated by the mirror neuron system and that it is a tightly controlled laboratory equivalent of the motor mimicry observed in naturalistic social contexts. If true, then automatic imitation can be used as a tool to investigate how the mirror neuron system contributes to cognitive functioning and how motor mimicry promotes prosocial attitudes and behavior.

Meta-analysis of imitation studies in humans suggest that there is enough evidence of mirror system activation during imitation that mirror neuron involvement is likely, even though no published studies have recorded the activities of singular neurons.