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Monday, July 29, 2019

Theory of mind

From Wikipedia, the free encyclopedia
 
Theory of mind is the ability to attribute mental states — beliefs, intents, desires, emotions, knowledge, etc. — to oneself, and to others, and to understand that others have beliefs, desires, intentions, and perspectives that are different from one's own. Theory of mind is crucial for everyday human social interactions and is used when analyzing, judging, and inferring others' behaviors. Deficits can occur in people with autism spectrum disorders, schizophrenia, attention deficit hyperactivity disorder, cocaine addiction, and brain damage suffered from alcohol's neurotoxicity. Although philosophical approaches to this exist, the theory of mind as such is distinct from the philosophy of mind.

Definition

Theory of mind is a theory insofar as the mind is the only thing being directly observed. The presumption that others have a mind is termed a theory of mind because each human can only intuit the existence of their own mind through introspection, and no one has direct access to the mind of another. It is typically assumed that others have minds analogous to one's own, and this assumption is based on the reciprocal, social interaction, as observed in joint attention, the functional use of language, and the understanding of others' emotions and actions. Having theory of mind allows one to attribute thoughts, desires, and intentions to others, to predict or explain their actions, and to posit their intentions. As originally defined, it enables one to understand that mental states can be the cause of—and thus be used to explain and predict—the behavior of others. Being able to attribute mental states to others and understanding them as causes of behavior implies, in part, that one must be able to conceive of the mind as a "generator of representations". If a person does not have a complete theory of mind, it may be a sign of cognitive or developmental impairment. 

Theory of mind appears to be an innate potential ability in humans that requires social and other experience over many years for its full development. Different people may develop more, or less, effective theory of mind. Neo-Piagetian theories of cognitive development maintain that theory of mind is a byproduct of a broader hypercognitive ability of the human mind to register, monitor, and represent its own functioning.

Empathy is a related concept, meaning the recognition and understanding of the states of mind of others, including their beliefs, desires and particularly emotions. This is often characterized as the ability to "put oneself into another's shoes". Recent neuro-ethological studies of animal behaviour suggest that even rodents may exhibit ethical or empathetic abilities. While empathy is known as emotional perspective-taking, theory of mind is defined as cognitive perspective-taking.

Research on theory of mind, in humans and animals, adults and children, normally and atypically developing, has grown rapidly in the 35 years since Premack and Guy Woodruff's paper, "Does the chimpanzee have a theory of mind?" The emerging field of social neuroscience has also begun to address this debate, by imaging the brains of humans while they perform tasks demanding the understanding of an intention, belief or other mental state in others. 

An alternative account of theory of mind is given within operant psychology and provides significant empirical evidence for a functional account of both perspective-taking and empathy. The most developed operant approach is founded on research on derived relational responding and is subsumed within what is called relational frame theory. According to this view, empathy and perspective-taking comprise a complex set of derived relational abilities based on learning to discriminate and respond verbally to ever more complex relations between self, others, place, and time, and through established relations.

Philosophical and psychological roots

Contemporary discussions of Theory of Mind have their roots in philosophical debate—most broadly, from the time of Descartes' Second Meditation, which set the groundwork for considering the science of the mind. Most prominent recently are two contrasting approaches in the philosophical literature, to theory of mind: theory-theory and simulation theory. The theory-theorist imagines a veritable theory—"folk psychology"—used to reason about others' minds. The theory is developed automatically and innately, though instantiated through social interactions. It is also closely related to person perception and attribution theory from social psychology. 

The intuitive assumption that others are minded is an apparent tendency we all share. We anthropomorphize non-human animals, inanimate objects, and even natural phenomena. Daniel Dennett referred to this tendency as taking an "intentional stance" toward things: we assume they have intentions, to help predict future behavior. However, there is an important distinction between taking an "intentional stance" toward something and entering a "shared world" with it. The intentional stance is a detached and functional theory we resort to during interpersonal interactions. A shared world is directly perceived and its existence structures reality itself for the perceiver. It is not just automatically applied to perception; it in many ways constitutes perception.

The philosophical roots of the relational frame theory (RFT) account of Theory of Mind arise from contextual psychology and refer to the study of organisms (both human and non-human) interacting in and with a historical and current situational context. It is an approach based on contextualism, a philosophy in which any event is interpreted as an ongoing act inseparable from its current and historical context and in which a radically functional approach to truth and meaning is adopted. As a variant of contextualism, RFT focuses on the construction of practical, scientific knowledge. This scientific form of contextual psychology is virtually synonymous with the philosophy of operant psychology.

Development

The study of which animals are capable of attributing knowledge and mental states to others, as well as the development of this ability in human ontogeny and phylogeny, has identified several behavioral precursors to theory of mind. Understanding attention, understanding of others' intentions, and imitative experience with other people are hallmarks of a theory of mind that may be observed early in the development of what later becomes a full-fledged theory. In studies with non-human animals and pre-verbal humans, in particular, researchers look to these behaviors preferentially in making inferences about mind. 

Simon Baron-Cohen identified the infant's understanding of attention in others, a social skill found by 7 to 9 months of age, as a "critical precursor" to the development of theory of mind. Understanding attention involves understanding that seeing can be directed selectively as attention, that the looker assesses the seen object as "of interest", and that seeing can induce beliefs. Attention can be directed and shared by the act of pointing, a joint attention behavior that requires taking into account another person's mental state, particularly whether the person notices an object or finds it of interest. Baron-Cohen speculates that the inclination to spontaneously reference an object in the world as of interest ("protodeclarative pointing") and to likewise appreciate the directed attention and interests of another may be the underlying motive behind all human communication.

Understanding of others' intentions is another critical precursor to understanding other minds because intentionality, or "aboutness", is a fundamental feature of mental states and events. The "intentional stance" has been defined by Daniel Dennett as an understanding that others' actions are goal-directed and arise from particular beliefs or desires. Both 2- and 3-year-old children could discriminate when an experimenter intentionally vs. accidentally marked a box with stickers as baited. Even earlier in ontogeny, Andrew N. Meltzoff found that 18-month-old infants could perform target manipulations that adult experimenters attempted and failed, suggesting the infants could represent the object-manipulating behavior of adults as involving goals and intentions. While attribution of intention (the box-marking) and knowledge (false-belief tasks) is investigated in young humans and nonhuman animals to detect precursors to a theory of mind, Gagliardi et al. have pointed out that even adult humans do not always act in a way consistent with an attributional perspective. In the experiment, adult human subjects made choices about baited containers when guided by confederates who could not see (and therefore, not know) which container was baited. 

Recent research in developmental psychology suggests that the infant's ability to imitate others lies at the origins of both theory of mind and other social-cognitive achievements like perspective-taking and empathy. According to Meltzoff, the infant's innate understanding that others are "like me" allows it to recognize the equivalence between the physical and mental states apparent in others and those felt by the self. For example, the infant uses his own experiences, orienting his head/eyes toward an object of interest to understand the movements of others who turn toward an object, that is, that they will generally attend to objects of interest or significance. Some researchers in comparative disciplines have hesitated to put a too-ponderous weight on imitation as a critical precursor to advanced human social-cognitive skills like mentalizing and empathizing, especially if true imitation is no longer employed by adults. A test of imitation by Alexandra Horowitz found that adult subjects imitated an experimenter demonstrating a novel task far less closely than children did. Horowitz points out that the precise psychological state underlying imitation is unclear and cannot, by itself, be used to draw conclusions about the mental states of humans. 

While much research has been done on infants, theory of mind develops continuously throughout childhood and into late adolescence as the synapses (neuronal connections) in the prefrontal cortex develop. The prefrontal cortex is thought to be involved in planning and decision-making. Children seem to develop theory of mind skills sequentially. The first skill to develop is the ability to recognize that others have diverse desires. Children are able to recognize that others have diverse beliefs soon after. The next skill to develop is recognizing that others have access to different knowledge bases. Finally, children are able to understand that others may have false beliefs and that others are capable of hiding emotions. While this sequence represents the general trend in skill acquisition, it seems that more emphasis is placed on some skills in certain cultures, leading to more valued skills to develop before those that are considered not as important. For example, in individualistic cultures such as the United States, a greater emphasis is placed on the ability to recognize that others have different opinions and beliefs. In a collectivistic culture, such as China, this skill may not be as important and therefore may not develop until later.

Language

There is evidence to believe that the development of theory of mind is closely intertwined with language development in humans. One meta-analysis showed a moderate to strong correlation (r = 0.43) between performance on theory of mind and language tasks. One might argue that this relationship is due solely to the fact that both language and theory of mind seem to begin to develop substantially around the same time in children (between ages 2–5). However, many other abilities develop during this same time period as well, and do not produce such high correlations with one another nor with theory of mind. There must be something else going on to explain the relationship between theory of mind and language. 

Pragmatic theories of communication assume that infants must possess an understanding of beliefs and mental states of others to infer the communicative content that proficient language users intend to convey. Since a verbal utterance is often underdetermined, and therefore, it can have different meanings depending on the actual context theory of mind abilities can play a crucial role in understanding the communicative and informative intentions of others and inferring the meaning of words. Some empirical results suggest that even 13-month-old infants have an early capacity for communicative mind-reading that enables them to infer what relevant information is transferred between communicative partners, which implies that human language relies at least partially on theory of mind skills.

Carol A. Miller posed further possible explanations for this relationship. One idea was that the extent of verbal communication and conversation involving children in a family could explain theory of mind development. The belief is that this type of language exposure could help introduce a child to the different mental states and perspectives of others. This has been suggested empirically by findings indicating that participation in family discussion predict scores on theory of mind tasks, as well as findings showing that deaf children who have hearing parents and may not be able to communicate with their parents much during early years of development tend to score lower on theory of mind tasks.

Another explanation of the relationship between language and theory of mind development has to do with a child's understanding of mental state words such as "think" and "believe". Since a mental state is not something that one can observe from behavior, children must learn the meanings of words denoting mental states from verbal explanations alone, requiring knowledge of the syntactic rules, semantic systems, and pragmatics of a language. Studies have shown that understanding of these mental state words predicts theory of mind in four-year-olds.

A third hypothesis is that the ability to distinguish a whole sentence ("Jimmy thinks the world is flat") from its embedded complement ("the world is flat") and understand that one can be true while the other can be false is related to theory of mind development. Recognizing these sentential complements as being independent of one another is a relatively complex syntactic skill and has been shown to be related to increased scores on theory of mind tasks in children.

In addition to these hypotheses, there is also evidence that the neural networks between the areas of the brain responsible for language and theory of mind are closely connected. The temporoparietal junction has been shown to be involved in the ability to acquire new vocabulary, as well as perceive and reproduce words. The temporoparietal junction also contains areas that specialize in recognizing faces, voices, and biological motion, in addition to theory of mind. Since all of these areas are located so closely together, it is reasonable to conclude that they work together. Moreover, studies have reported an increase in activity in the TPJ when patients are absorbing information through reading or images regarding other peoples' beliefs but not while observing information about physical control stimuli.

Theory of mind in adults

Neurotypical adults have the theory of mind concepts that they developed as children (concepts such as belief, desire, knowledge and intention). A focal question is how they use these concepts to meet the diverse demands of social life, ranging from snap decisions about how to trick an opponent in a competitive game, to keeping up with who knows what in a fast-moving conversation, to judging the guilt or innocence of the accused in a court of law.

Boaz Keysar, Dale Barr and colleagues found that adults often failed to use their theory of mind abilities to interpret a speaker’s message, even though they were perfectly well aware that the speaker lacked critical knowledge. Other studies converge in showing that adults are prone to “egocentric biases”, whereby they are influenced by their own beliefs, knowledge or preferences when judging those of other people, or else neglect other people’s perspectives entirely. There is also evidence that adults with greater memory and inhibitory capacity and greater motivation are more likely to use their theory of mind abilities.

In contrast, evidence from tasks looking for indirect effects of thinking about other people’s mental states suggests that adults may sometimes use their theory of mind automatically. Agnes Kovacs and colleagues measured the time it took adults to detect the presence of a ball as it was revealed from behind an occluder. They found that adults’ speed of response was influenced by whether or not an avatar in the scene thought there was a ball behind the occluder, even though adults were not asked to pay attention to what the avatar thought. Dana Samson and colleagues measured the time it took adults to judge the number of dots on the wall of a room. They found that adults responded more slowly when an avatar standing in the room happened to see fewer dots than they did, even when they had never been asked to pay attention to what the avatar could see. It has been questioned whether these “altercentric biases” truly reflect automatic processing of what another person is thinking or seeing, or whether they instead reflect attention and memory effects cued by the avatar, but not involving any representation of what they think or see.

Different theories have sought to explain these patterns of results. The idea that theory of mind is automatic is attractive because it would help explain how people keep up with the theory of mind demands of competitive games and fast-moving conversations. It might also explain evidence that human infants and some non-human species sometimes appear capable of theory of mind, despite their limited resources for memory and cognitive control. The idea that theory of mind is effortful and not automatic is attractive because it feels effortful to decide whether a defendant is guilty or innocent, or whether a negotiator is bluffing, and economy of effort would help explain why people sometimes neglect to use their theory of mind. Ian Apperly and Stephen Butterfill have suggested that people do in fact have “two systems” for theory of mind, in common with “two systems” accounts in many other areas of psychology. On this account, “system 1” is cognitively efficient and enables theory of mind for a limited but useful set of circumstances. “System 2” is cognitively effortful, but enables much more flexible theory of mind abilities. This account has been criticised by Peter Carruthers who suggests that the same core theory of mind abilities can be used in both simple and complex ways. The account has been criticised by Celia Heyes who suggests that “system 1” theory of mind abilities do not require representation of mental states of other people, and so are better thought of as “sub-mentalising”.

Aging

In older age, theory of mind capacities decline, irrespective of how exactly they are tested (e.g. stories, eyes, videos, false belief-video, false belief-other, faux pas). However, the decline in other cognitive functions is even stronger, suggesting that social cognition is somewhat preserved. In contrast to theory of mind, empathy shows no impairments in aging.

There are two kinds of theory of mind representations: cognitive (concerning the mental states, beliefs, thoughts, and intentions of others) and affective (concerning the emotions of others). Cognitive theory of mind is further separated into first order (e.g., I think she thinks that…) and second order (e.g., he thinks that she thinks that…). There is evidence that cognitive and affective theory of mind processes are functionally independent from one another. In studies of Alzheimer’s disease, which typically occurs in older adults, the patients display impairment with second order cognitive theory of mind, but usually not with first order cognitive or affective theory of mind. However, it is difficult to discern a clear pattern of theory of mind variation due to age. There have been many discrepancies in the data collected thus far, likely due to small sample sizes and the use of different tasks that only explore one aspect of theory of mind. Many researchers suggest that the theory of mind impairment is simply due to the normal decline in cognitive function.

Cultural variations

Researchers have proposed that five key aspects of theory of mind develop sequentially for all children between the ages of three to five. This five-step theory of mind scale consists of the development of diverse desires (DD), diverse beliefs (DB), knowledge access (KA), false beliefs (FB), and hidden emotions (HE). Australian, American and European children acquire theory of mind in this exact order, and studies with children in Canada, India, Peru, Samoa, and Thailand indicate that they all pass the false belief task at around the same time, suggesting that the children develop theory of mind consistently around the world.

However, children from Iran and China develop theory of mind in a slightly different order. Although they begin the development of theory of mind around the same time, toddlers from these countries understand knowledge access (KA) before Western children but take longer to understand false beliefs (FB). Researchers believe this swap in the developmental order is related to the culture of collectivism in Iran and China, which emphasizes interdependence and shared knowledge as opposed to the culture of individualism in Western countries, which promotes individuality and conflicting opinions. Because of these different cultural values, Iranian and Chinese children might take longer to understand that other people have different, sometimes false, beliefs. This suggests that the development of theory of mind is not universal and solely determined by innate brain processes but also influenced by social and cultural factors.

Empirical investigation

Whether children younger than 3 or 4 years old may have any theory of mind is a topic of debate among researchers. It is a challenging question, due to the difficulty of assessing what pre-linguistic children understand about others and the world. Tasks used in research into the development of Theory of Mind must take into account the umwelt—(the German word Umwelt means "environment" or "surrounding world")—of the pre-verbal child.

False-belief task

One of the most important milestones in theory of mind development is gaining the ability to attribute false belief: that is, to recognize that others can have beliefs about the world that are diverging. To do this, it is suggested, one must understand how knowledge is formed, that people's beliefs are based on their knowledge, that mental states can differ from reality, and that people's behavior can be predicted by their mental states. Numerous versions of the false-belief task have been developed, based on the initial task done by Wimmer and Perner (1983).

In the most common version of the false-belief task (often called the "'Sally-Anne' test" or "'Sally-Anne' task"), children are told or shown a story involving two characters. For example, the child is shown two dolls, Sally and Anne, who have a basket and a box, respectively. Sally also has a marble, which she places into her basket, and then leaves the room. While she is out of the room, Anne takes the marble from the basket and puts it into the box. Sally returns, and the child is then asked where Sally will look for the marble. The child passes the task if she answers that Sally will look in the basket, where Sally put the marble; the child fails the task if she answers that Sally will look in the box, where the child knows the marble is hidden, even though Sally cannot know this, since she did not see it hidden there. To pass the task, the child must be able to understand that another's mental representation of the situation is different from their own, and the child must be able to predict behavior based on that understanding.

Another example is when a boy leaves chocolate on a shelf and then leaves the room. His mother puts it in the fridge. To pass the task, the child must understand that the boy, upon returning, holds the false belief that his chocolate is still on the shelf.

The results of research using false-belief tasks have been fairly consistent: most normally developing children are able to pass the tasks from around age four. Notably, while most children, including those with Down syndrome, are able to pass this test, in one study, 80% of children diagnosed with autism were unable to do so.

Also adults can experience problems with false beliefs. For instance, when they show hindsight bias, defined as: "the inclination to see events that have already happened as being more predictable than they were before they took place." In an experiment by Fischhoff in 1975, adult subjects who were asked for an independent assessment were unable to disregard information on actual outcome. Also in experiments with complicated situations, when assessing others' thinking, adults can be unable to disregard certain information that they have been given.

Unexpected contents

Other tasks have been developed to try to solve the problems inherent in the false-belief task. In the "Unexpected contents", or "Smarties" task, experimenters ask children what they believe to be the contents of a box that looks as though it holds a candy called "Smarties". After the child guesses (usually) "Smarties", it is shown that the box in fact contained pencils. The experimenter then re-closes the box and asks the child what she thinks another person, who has not been shown the true contents of the box, will think is inside. The child passes the task if he/she responds that another person will think that "Smarties" exist in the box, but fails the task if she responds that another person will think that the box contains pencils. Gopnik & Astington (1988) found that children pass this test at age four or five years.

Other tasks

The "false-photograph" task is another task that serves as a measure of theory of mind development. In this task, children must reason about what is represented in a photograph that differs from the current state of affairs. Within the false-photograph task, either a location or identity change exists. In the location-change task, the examiner puts an object in one location (e.g., chocolate in an open green cupboard), whereupon the child takes a Polaroid photograph of the scene. While the photograph is developing, the examiner moves the object to a different location (e.g., a blue cupboard), allowing the child to view the examiner's action. The examiner asks the child two control questions: "When we first took the picture, where was the object?" and "Where is the object now?". The subject is also asked a "false-photograph" question: "Where is the object in the picture?" The child passes the task if he/she correctly identifies the location of the object in the picture and the actual location of the object at the time of the question. However, the last question might be misinterpreted as: "Where in this room is the object that the picture depicts?" and therefore some examiners use an alternative phrasing.

To make it easier for animals, young children, and individuals with classical (Kanner-type) autism to understand and perform theory of mind tasks, researchers have developed tests in which verbal communication is de-emphasized: some whose administration does not involve verbal communication on the part of the examiner, some whose successful completion does not require verbal communication on the part of the subject, and some that meet both of the foregoing standards. One category of tasks uses a preferential looking paradigm, with looking time as the dependent variable. For instance, 9-month-old infants prefer looking at behaviors performed by a human hand over those made by an inanimate hand-like object. Other paradigms look at rates of imitative behavior, the ability to replicate and complete unfinished goal-directed acts, and rates of pretend play.

Early precursors

Recent research on the early precursors of theory of mind has looked at innovative ways at capturing preverbal infants' understanding of other people's mental states, including perception and beliefs. Using a variety of experimental procedures, studies have shown that infants from their first year of life have an implicit understanding of what other people see and what they know. A popular paradigm used to study infants' theory of mind is the violation of expectation procedure, which predicates on infants' tendency to look longer at unexpected and surprising events compared to familiar and expected events. Therefore, their looking-times measures would give researchers an indication of what infants might be inferring, or their implicit understanding of events. One recent study using this paradigm found that 16-month-olds tend to attribute beliefs to a person whose visual perception was previously witnessed as being "reliable", compared to someone whose visual perception was "unreliable". Specifically, 16-month-olds were trained to expect a person's excited vocalization and gaze into a container to be associated with finding a toy in the reliable-looker condition or an absence of a toy in the unreliable-looker condition. Following this training phase, infants witnessed, in an object-search task, the same persons either searching for a toy in the correct or incorrect location after they both witnessed the location of where the toy was hidden. Infants who experienced the reliable looker were surprised and therefore looked longer when the person searched for the toy in the incorrect location compared to the correct location. In contrast, the looking time for infants who experienced the unreliable looker did not differ for either search locations. These findings suggest that 16-month-old infants can differentially attribute beliefs about a toy's location based on the person's prior record of visual perception.

Deficits

The theory of mind impairment describes a difficulty someone would have with perspective-taking. This is also sometimes referred to as mind-blindness. This means that individuals with a theory of mind impairment would have a difficult time seeing phenomena from any other perspective than their own. Individuals who experience a theory of mind deficit have difficulty determining the intentions of others, lack understanding of how their behavior affects others, and have a difficult time with social reciprocity. Theory of Mind deficits have been observed in people with autism spectrum disorders, people with schizophrenia, people with nonverbal learning disorder, people with attention deficit disorder, persons under the influence of alcohol and narcotics, sleep-deprived persons, and persons who are experiencing severe emotional or physical pain. Theory of mind deficits have also been observed in deaf children who are late signers (i.e., are born to hearing parents), but the deficit is due to the delay in language learning, not any cognitive deficit, and therefore disappears once the child learns sign language.

Autism

In 1985 Simon Baron-Cohen, Alan M. Leslie and Uta Frith suggested that children with autism do not employ theory of mind and suggested that autistic children have particular difficulties with tasks requiring the child to understand another person's beliefs. These difficulties persist when children are matched for verbal skills and have been taken as a key feature of autism. 

Many individuals classified as autistic have severe difficulty assigning mental states to others, and they seem to lack theory of mind capabilities. Researchers who study the relationship between autism and theory of mind attempt to explain the connection in a variety of ways. One account assumes that theory of mind plays a role in the attribution of mental states to others and in childhood pretend play. According to Leslie, theory of mind is the capacity to mentally represent thoughts, beliefs, and desires, regardless of whether or not the circumstances involved are real. This might explain why some autistic individuals show extreme deficits in both theory of mind and pretend play. However, Hobson proposes a social-affective justification, which suggests that with an autistic person, deficits in theory of mind result from a distortion in understanding and responding to emotions. He suggests that typically developing human beings, unlike autistic individuals, are born with a set of skills (such as social referencing ability) that later lets them comprehend and react to other people's feelings. Other scholars emphasize that autism involves a specific developmental delay, so that autistic children vary in their deficiencies, because they experience difficulty in different stages of growth. Very early setbacks can alter proper advancement of joint-attention behaviors, which may lead to a failure to form a full theory of mind.

It has been speculated that Theory of Mind exists on a continuum as opposed to the traditional view of a discrete presence or absence. While some research has suggested that some autistic populations are unable to attribute mental states to others, recent evidence points to the possibility of coping mechanisms that facilitate a spectrum of mindful behavior. Tine et al. suggest that autistic children score substantially lower on measures of social theory of mind in comparison to children diagnosed with Asperger syndrome.

Generally, children with more advanced theory of mind abilities display more advanced social skills, greater adaptability to new situations, and greater cooperation with others. As a result, these children are typically well-liked. However, “children may use their mind-reading abilities to manipulate, outwit, tease, or trick their peers”. Individuals possessing inferior theory of mind skills, such as children with autism spectrum disorder, may be socially rejected by their peers since they are unable to communicate effectively. Social rejection has been proven to negatively impact a child’s development and can put the child at greater risk of developing depressive symptoms.

Peer-mediated interventions (PMI) are a school-based treatment approach for children and adolescents with autism spectrum disorder in which peers are trained to be role models in order to promote social behavior. Laghi et al. studied if analysis of prosocial (nice) and antisocial (nasty) theory of mind behaviors could be used, in addition to teacher recommendations, to select appropriate candidates for PMI programs. Selecting children with advanced theory of mind skills who use them in prosocial ways will theoretically make the program more effective. While the results indicated that analyzing the social uses of theory of mind of possible candidates for a PMI program is invaluable, it may not be a good predictor of a candidate's performance as a role model.

Schizophrenia

Individuals with the diagnosis of schizophrenia can show deficits in theory of mind. Mirjam Sprong and colleagues investigated the impairment by examining 29 different studies, with a total of over 1500 participants. This meta-analysis showed significant and stable deficit of theory of mind in people with schizophrenia. They performed poorly on false-belief tasks, which test the ability to understand that others can hold false beliefs about events in the world, and also on intention-inference tasks, which assess the ability to infer a character's intention from reading a short story. Schizophrenia patients with negative symptoms, such as lack of emotion, motivation, or speech, have the most impairment in theory of mind and are unable to represent the mental states of themselves and of others. Paranoid schizophrenic patients also perform poorly because they have difficulty accurately interpreting others' intentions. The meta-analysis additionally showed that IQ, gender, and age of the participants does not significantly affect the performance of theory of mind tasks.

Current research suggests that impairment in theory of mind negatively affects clinical insight, the patient's awareness of their mental illness. Insight requires theory of mind—a patient must be able to adopt a third-person perspective and see the self as others do. A patient with good insight would be able to accurately self-represent, by comparing oneself with others and by viewing oneself from the perspective of others. Insight allows a patient to recognize and react appropriately to his symptoms; however, a patient who lacks insight would not realize that he has a mental illness, because of his inability to accurately self-represent. Therapies that teach patients perspective-taking and self-reflection skills can improve abilities in reading social cues and taking the perspective of another person.

The majority of the current literature supports the argument that the theory of mind deficit is a stable trait-characteristic rather than a state-characteristic of schizophrenia. The meta-analysis conducted by Sprong et al. showed that patients in remission still had impairment in theory of mind. The results indicate that the deficit is not merely a consequence of the active phase of schizophrenia.

Schizophrenic patients' deficit in theory of mind impairs their daily interactions with others. An example of a disrupted interaction is one between a schizophrenic parent and a child. Theory of mind is particularly important for parents, who must understand the thoughts and behaviors of their children and react accordingly. Dysfunctional parenting is associated with deficits in the first-order theory of mind, the ability to understand another person's thoughts, and the second-order theory of mind, the ability to infer what one person thinks about another person's thoughts. Compared with healthy mothers, mothers with schizophrenia are found to be more remote, quiet, self-absorbed, insensitive, unresponsive, and to have fewer satisfying interactions with their children. They also tend to misinterpret their children's emotional cues, and often misunderstand neutral faces as negative. Activities such as role-playing and individual or group-based sessions are effective interventions that help the parents improve on perspective-taking and theory of mind. Although there is a strong association between theory of mind deficit and parental role dysfunction, future studies could strengthen the relationship by possibly establishing a causal role of theory of mind on parenting abilities.

Alcohol use disorders

Impairments in theory of mind, as well as other social-cognitive deficits are commonly found in people suffering from alcoholism, due to the neurotoxic effects of alcohol on the brain, particularly the prefrontal cortex.

Depression and dysphoria

Individuals in a current major depressive episode, a disorder characterized by social impairment, show deficits in theory of mind decoding. Theory of mind decoding is the ability to use information available in the immediate environment (e.g., facial expression, tone of voice, body posture) to accurately label the mental states of others. The opposite pattern, enhanced theory of mind, is observed in individuals vulnerable to depression, including those individuals with past major depressive disorder (MDD), dysphoric individuals, and individuals with a maternal history of MDD.

Developmental language disorder

Children diagnosed with developmental language disorder (DLD) exhibit much lower scores on reading and writing sections of standardized tests, yet have a normal nonverbal IQ. These language deficits can be any specific deficits in lexical semantics, syntax, or pragmatics, or a combination of multiple problems. They often exhibit poorer social skills than normally developing children, and seem to have problems decoding beliefs in others. A recent meta-analysis confirmed that children with DLD have substantially lower scores on theory of mind tasks compared to typically developing children.  This strengthens the claim that language development is related to theory of mind.

Brain mechanisms

In typically developing humans

Research on theory of mind in autism led to the view that mentalizing abilities are subserved by dedicated mechanisms that can - in some cases - be impaired while general cognitive function remains largely intact. 

Neuroimaging research has supported this view, demonstrating specific brain regions consistently engaged during theory of mind tasks. PET research on theory of mind, using verbal and pictorial story comprehension tasks, has identified a set of brain regions including the medial prefrontal cortex (mPFC), and area around posterior superior temporal sulcus (pSTS), and sometimes precuneus and amygdala/temporopolar cortex. Subsequently, research on the neural basis of theory of mind has diversified, with separate lines of research focused on the understanding of beliefs, intentions, and more complex properties of minds such as psychological traits.

Studies from Rebecca Saxe's lab at MIT, using a false-belief versus false-photograph task contrast aimed at isolating the mentalizing component of the false-belief task, have very consistently found activation in mPFC, precuneus, and temporo-parietal junction (TPJ), right-lateralized. In particular, it has been proposed that the right TPJ (rTPJ) is selectively involved in representing the beliefs of others. However, some debate exists, as some scientists have noted that the same rTPJ region has been consistently activated during spatial reorienting of visual attention; Jean Decety from the University of Chicago and Jason Mitchell from Harvard have thus proposed that the rTPJ subserves a more general function involved in both false-belief understanding and attentional reorienting, rather than a mechanism specialized for social cognition. However, it is possible that the observation of overlapping regions for representing beliefs and attentional reorienting may simply be due to adjacent, but distinct, neuronal populations that code for each. The resolution of typical fMRI studies may not be good enough to show that distinct/adjacent neuronal populations code for each of these processes. In a study following Decety and Mitchell, Saxe and colleagues used higher-resolution fMRI and showed that the peak of activation for attentional reorienting is approximately 6-10mm above the peak for representing beliefs. Further corroborating that differing populations of neurons may code for each process, they found no similarity in the patterning of fMRI response across space.

Functional imaging has also been used to study the detection of mental state information in Heider-Simmel-esque animations of moving geometric shapes, which typical humans automatically perceive as social interactions laden with intention and emotion. Three studies found remarkably similar patterns of activation during the perception of such animations versus a random or deterministic motion control: mPFC, pSTS, fusiform face area (FFA), and amygdala were selectively engaged during the Theory of Mind condition. Another study presented subjects with an animation of two dots moving with a parameterized degree of intentionality (quantifying the extent to which the dots chased each other), and found that pSTS activation correlated with this parameter.

A separate body of research has implicated the posterior superior temporal sulcus in the perception of intentionality in human action; this area is also involved in perceiving biological motion, including body, eye, mouth, and point-light display motion. One study found increased pSTS activation while watching a human lift his hand versus having his hand pushed up by a piston (intentional versus unintentional action). Several studies have found increased pSTS activation when subjects perceive a human action that is incongruent with the action expected from the actor's context and inferred intention. Examples would be: a human performing a reach-to-grasp motion on empty space next to an object, versus grasping the object; a human shifting eye gaze toward empty space next to a checkerboard target versus shifting gaze toward the target; an unladen human turning on a light with his knee, versus turning on a light with his knee while carrying a pile of books; and a walking human pausing as he passes behind a bookshelf, versus walking at a constant speed. In these studies, actions in the "congruent" case have a straightforward goal, and are easy to explain in terms of the actor's intention. The incongruent actions, on the other hand, require further explanation (why would someone twist empty space next to a gear?), and then apparently would demand more processing in the STS. Note that this region is distinct from the temporo-parietal area activated during false belief tasks. Also note that pSTS activation in most of the above studies was largely right-lateralized, following the general trend in neuroimaging studies of social cognition and perception. Also right-lateralized are the TPJ activation during false belief tasks, the STS response to biological motion, and the FFA response to faces. 

Neuropsychological evidence has provided support for neuroimaging results regarding the neural basis of theory of mind. Studies with patients suffering from a lesion of the frontal lobes and the temporoparietal junction of the brain (between the temporal lobe and parietal lobe) reported that they have difficulty with some theory of mind tasks. This shows that theory of mind abilities are associated with specific parts of the human brain. However, the fact that the medial prefrontal cortex and temporoparietal junction are necessary for theory of mind tasks does not imply that these regions are specific to that function. TPJ and mPFC may subserve more general functions necessary for Theory of Mind. 

Research by Vittorio Gallese, Luciano Fadiga and Giacomo Rizzolatti (reviewed in) has shown that some sensorimotor neurons, which are referred to as mirror neurons, first discovered in the premotor cortex of rhesus monkeys, may be involved in action understanding. Single-electrode recording revealed that these neurons fired when a monkey performed an action, as well as when the monkey viewed another agent carrying out the same task. Similarly, fMRI studies with human participants have shown brain regions (assumed to contain mirror neurons) that are active when one person sees another person's goal-directed action. These data have led some authors to suggest that mirror neurons may provide the basis for theory of mind in the brain, and to support simulation theory of mind reading (see above).

There is also evidence against the link between mirror neurons and theory of mind. First, macaque monkeys have mirror neurons but do not seem to have a 'human-like' capacity to understand theory of mind and belief. Second, fMRI studies of theory of mind typically report activation in the mPFC, temporal poles and TPJ or STS, but these brain areas are not part of the mirror neuron system. Some investigators, like developmental psychologist Andrew Meltzoff and neuroscientist Jean Decety, believe that mirror neurons merely facilitate learning through imitation and may provide a precursor to the development of Theory of Mind. Others, like philosopher Shaun Gallagher, suggest that mirror-neuron activation, on a number of counts, fails to meet the definition of simulation as proposed by the simulation theory of mindreading.

In a recent paper, Keren Haroush and Ziv Williams outlined the case for a group of neurons in primates' brains that uniquely predicted the choice selection of their interacting partner. These primates' neurons, located in the anterior cingulate cortex of rhesus monkeys, were observed using single-unit recording while the monkeys played a variant of the iterative prisoner's dilemma game. By identifying cells that represent the yet unknown intentions of a game partner, Haroush & Williams' study supports the idea that theory of mind may be a fundamental and generalized process, and suggests that anterior cingulate cortex neurons may potentially act to complement the function of mirror neurons during social interchange.

In autism

Several neuroimaging studies have looked at the neural basis theory of mind impairment in subjects with Asperger syndrome and high-functioning autism (HFA). The first PET study of theory of mind in autism (also the first neuroimaging study using a task-induced activation paradigm in autism) replicated a prior study in normal individuals, which employed a story-comprehension task. This study found displaced and diminished mPFC activation in subjects with autism. However, because the study used only six subjects with autism, and because the spatial resolution of PET imaging is relatively poor, these results should be considered preliminary.

A subsequent fMRI study scanned normally developing adults and adults with HFA while performing a "reading the mind in the eyes" task: viewing a photo of a human's eyes and choosing which of two adjectives better describes the person's mental state, versus a gender discrimination control. The authors found activity in orbitofrontal cortex, STS, and amygdala in normal subjects, and found no amygdala activation and abnormal STS activation in subjects with autism. 

A more recent PET study looked at brain activity in individuals with HFA and Asperger syndrome while viewing Heider-Simmel animations (see above) versus a random motion control. In contrast to normally developing subjects, those with autism showed no STS or FFA activation, and significantly less mPFC and amygdala activation. Activity in extrastriate regions V3 and LO was identical across the two groups, suggesting intact lower-level visual processing in the subjects with autism. The study also reported significantly less functional connectivity between STS and V3 in the autism group. Note, however, that decreased temporal correlation between activity in STS and V3 would be expected simply from the lack of an evoked response in STS to intent-laden animations in subjects with autism. A more informative analysis would be to compute functional connectivity after regressing out evoked responses from all-time series.

A subsequent study, using the incongruent/congruent gaze-shift paradigm described above, found that in high-functioning adults with autism, posterior STS (pSTS) activation was undifferentiated while they watched a human shift gaze toward a target and then toward adjacent empty space. The lack of additional STS processing in the incongruent state may suggest that these subjects fail to form an expectation of what the actor should do given contextual information, or that feedback about the violation of this expectation doesn't reach STS. Both explanations involve an impairment in the ability to link eye gaze shifts with intentional explanations. This study also found a significant anticorrelation between STS activation in the incongruent-congruent contrast and social subscale score on the Autism Diagnostic Interview-Revised, but not scores on the other subscales. 

In 2011, an fMRI study demonstrated that the right temporoparietal junction (rTPJ) of higher-functioning adults with autism was not more selectively activated for mentalizing judgments when compared to physical judgments about self and other. rTPJ selectivity for mentalizing was also related to individual variation on clinical measures of social impairment: individuals whose rTPJ was increasingly more active for mentalizing compared to physical judgments were less socially impaired, while those who showed little to no difference in response to mentalizing or physical judgments were the most socially impaired. This evidence builds on work in typical development that suggests rTPJ is critical for representing mental state information, irrespective of whether it is about oneself or others. It also points to an explanation at the neural level for the pervasive mind-blindness difficulties in autism that are evident throughout the lifespan.

In schizophrenia

The brain regions associated with theory of mind include the superior temporal gyrus (STS), the temporoparietal junction (TPJ), the medial prefrontal cortex (MPFC), the precuneus, and the amygdala. The reduced activity in the MPFC of individuals with schizophrenia is associated with the Theory of mind deficit and may explain impairments in social function among people with schizophrenia. Increased neural activity in MPFC is related to better perspective-taking, emotion management, and increased social functioning. Disrupted brain activities in areas related to theory of mind may increase social stress or disinterest in social interaction, and contribute to the social dysfunction associated with schizophrenia.

Practical validity

Group member average scores of theory of mind abilities, measured with the Reading the Mind in the Eyes test (RME), are suggested as drivers of successful group performance. In particular, high group average scores on the RME are shown to be correlated with the collective intelligence factor c defined as a group's ability to perform a wide range of mental tasks, a group intelligence measure similar to the g factor for general individual intelligence. RME is a Theory of Mind test for adults that shows sufficient test-retest reliability and constantly differentiates control groups from individuals with functional autism or Asperger syndrome. It is one of the most widely accepted and well-validated tests for Theory of Mind abilities within adults.

Evolution

The evolutionary origin of theory of mind remains obscure. While many theories make claims about its role in the development of human language and social cognition few of them specify in detail any evolutionary neurophysiological precursors. A recent theory claims that Theory of Mind has its roots in two defensive reactions, namely immobilization stress and tonic immobility, which are implicated in the handling of stressful encounters and also figure prominently in mammalian childrearing practices (Tsoukalas, 2018). Their combined effect seems capable of producing many of the hallmarks of theory of mind, e.g., eye-contact, gaze-following, inhibitory control and intentional attributions.

Non-human

An open question is whether other animals besides humans have a genetic endowment and social environment that allows them to acquire a theory of mind in the same way that human children do. This is a contentious issue because of the problem of inferring from animal behavior the existence of thinking or of particular thoughts, or the existence of a concept of self or self-awareness, consciousness and qualia. One difficulty with non-human studies of theory of mind is the lack of sufficient numbers of naturalistic observations, giving insight into what the evolutionary pressures might be on a species' development of theory of mind. 

Non-human research still has a major place in this field, however, and is especially useful in illuminating which nonverbal behaviors signify components of theory of mind, and in pointing to possible stepping points in the evolution of what many claim to be a uniquely human aspect of social cognition. While it is difficult to study human-like theory of mind and mental states in species whose potential mental states we have an incomplete understanding, researchers can focus on simpler components of more complex capabilities. For example, many researchers focus on animals' understanding of intention, gaze, perspective, or knowledge (or rather, what another being has seen). A study that looked at understanding of intention in orangutans, chimpanzees and children showed that all three species understood the difference between accidental and intentional acts. Part of the difficulty in this line of research is that observed phenomena can often be explained as simple stimulus-response learning, as it is in the nature of any theorizers of mind to have to extrapolate internal mental states from observable behavior. Recently, most non-human theory of mind research has focused on monkeys and great apes, who are of most interest in the study of the evolution of human social cognition. Other studies relevant to attributions theory of mind have been conducted using plovers and dogs, and have shown preliminary evidence of understanding attention—one precursor of theory of mind—in others. 

There has been some controversy over the interpretation of evidence purporting to show theory of mind ability—or inability—in animals. Two examples serve as demonstration: first, Povinelli et al. (1990) presented chimpanzees with the choice of two experimenters from whom to request food: one who had seen where food was hidden, and one who, by virtue of one of a variety of mechanisms (having a bucket or bag over his head; a blindfold over his eyes; or being turned away from the baiting) does not know, and can only guess. They found that the animals failed in most cases to differentially request food from the "knower". By contrast, Hare, Call, and Tomasello (2001) found that subordinate chimpanzees were able to use the knowledge state of dominant rival chimpanzees to determine which container of hidden food they approached. William Field and Sue Savage-Rumbaugh believe that bonobos have developed theory of mind, and cite their communications with a captive bonobo, Kanzi, as evidence.

In a 2016 experiment, ravens Corvus corax were shown to take into account visual access of unseen conspecifics. The researchers argued that "ravens can generalize from their own perceptual experience to infer the possibility of being seen".

A 2016 study published by evolutionary anthropologist Christopher Krupenye brings new light to the existence of Theory of Mind, and particularly false beliefs, in non-human primates.

Neuroscience and intelligence

From Wikipedia, the free encyclopedia
Neuroscience and intelligence refers to the various neurological factors that are partly responsible for the variation of intelligence within a species or between different species. A large amount of research in this area has been focused on the neural basis of human intelligence. Historic approaches to study the neuroscience of intelligence consisted of correlating external head parameters, for example head circumference, to intelligence. Post-mortem measures of brain weight and brain volume have also been used. More recent methodologies focus on examining correlates of intelligence within the living brain using techniques such as magnetic resonance imaging (MRI), functional MRI (fMRI), electroencephalography (EEG), positron emission tomography and other non-invasive measures of brain structure and activity.

Researchers have been able to identify correlates of intelligence within the brain and its functioning. These include overall brain volume, grey matter volume, white matter volume, white matter integrity, cortical thickness and neural efficiency. Although the evidence base for our understanding of the neural basis of human intelligence has increased greatly over the past 30 years, even more research is needed to fully understand it.

The neural basis of intelligence has also been examined in animals such as primates, cetaceans and rodents.

Humans

Brain volume

One of the main methods used to establish a relationship between intelligence and the brain is to use measures of Brain volume. The earliest attempts at estimating brain volume were done using measures of external head parameters, such as head circumference as a proxy for brain size. More recent methodologies employed to study this relationship include post-mortem measures of brain weight and volume. These have their own limitations and strengths. The advent of MRI as a non-invasive highly-accurate measure of living brain structure and function (using fMRI) made this the pre-dominant and preferred method for measuring brain volume.

Overall, larger brain size and volume is associated with better cognitive functioning and higher intelligence. The specific regions that show the most robust correlation between volume and intelligence are the frontal, temporal and parietal lobes of the brain. A large number of studies have been conducted with uniformly positive correlations, leading to the generally safe conclusion that larger brains predict greater intelligence. In healthy adults, the correlation of total brain volume and IQ is ~ 0.4 

Less is known about variation on scales less than total brain volume. A meta-analytic review by McDaniel found that the correlation between Intelligence and in vivo brain size was larger for females (0.40) than for males (0.25). The same study also found that the correlation between brain size and Intelligence increased with age, with children showing smaller correlations. It has been suggested that the link between larger brain volumes and higher intelligence is related to variation in specific brain regions: a whole-brain measure would under-estimate these links. For functions more specific than general intelligence, regional effects may be more important. For instance evidence suggests that in adolescents learning new words, vocabulary growth is associated with gray matter density in bilateral posterior supramarginal gyri. Small studies have shown transient changes in gray-matter associated with developing a new physical skill (juggling) occipito-temporal cortex.
 
Brain volume is not a perfect account of intelligence: the relationship explains a modest amount of variance in intelligence – 12% to 36% of the variance. The amount of variance explained by brain volume may also depend on the type of intelligence measured. Up to 36% of variance in verbal intelligence can be explained by brain volume, while only approximately 10% of variance in visuospatial intelligence can be explained by brain volume. A 2015 study by researcher Stuart J. Ritchie found that brain size explained 12% of the variance in intelligence among individuals. These caveats imply that there are other major factors influencing how intelligent an individual is apart from brain size. In a large meta-analysis consisting of 88 studies Pietschnig et al. (2015) estimated the correlation between brain volume and intelligence to be about correlation coefficient of 0.24 which equates to 6% variance. Taking into account measurement quality, and sample type and IQ-range, the meta-analytic association of brain volume in appears to be ~ .4 in normal adults. Researcher Jakob Pietschnig argued that the strength of the positive association of brain volume and IQ remains robust, but has been overestimated in the literature. He has stated that "It is tempting to interpret this association in the context of human cognitive evolution and species differences in brain size and cognitive ability, we show that it is not warranted to interpret brain size as an isomorphic proxy of human intelligence differences".

Grey matter

Grey matter has been examined as a potential biological foundation for differences in intelligence. Similarly to brain volume, global grey matter volume is positively associated with intelligence. More specifically, higher intelligence has been associated with larger cortical grey matter in the prefrontal and posterior temporal cortex in adults. Furthermore, both verbal and nonverbal intelligence have been shown to be positively correlated with grey matter volume across the parietal, temporal and occipital lobes in young healthy adults, implying that intelligence is associated with a wide variety of structures within the brain.

There appear to be sex differences between the relationship of grey matter to intelligence between men and women. Men appear to show more intelligence to grey matter correlations in the frontal and parietal lobes, while the strongest correlations between intelligence and grey matter in women can be found in the frontal lobes and Broca's area. However, these differences do not seem to impact overall Intelligence, implying that the same cognitive ability levels can be attained in different ways.

One specific methodology used to study grey matter correlates of intelligence in areas of the brain is known as voxel-based morphometry (VBM). VBM allows researchers to specify areas of interest with great spatial resolution, allowing the examination of grey matter areas correlated with intelligence with greater special resolution. VBM has been used to correlate grey matter positively with intelligence in the frontal, temporal, parietal, and occipital lobes in healthy adults. VBM has also been used to show that grey matter volume in the medial region of the prefrontal cortex and the dorsomedial prefrontal cortex correlate positively with intelligence in a group of 55 healthy adults. VBM has also been successfully used to establish a positive correlation between grey matter volumes in the anterior cingulate and intelligence in children aged 5 to 18 years old.

Grey matter has also been shown to positively correlate with intelligence in children. Reis and colleagues have found that grey matter in the prefrontal cortex contributes most robustly to variance in Intelligence in children between 5 and 17, while subcortical grey matter is related to intelligence to a lesser extent. Frangou and colleagues examined the relationship between grey matter and intelligence in children and young adults aged between 12 and 21, and found that grey matter in the orbitofrontal cortex, cingulate gyrus, cerebellum and thalamus was positively correlated to intelligence, while grey matter in the caudate nucleus is negatively correlated with intelligence. However, the relationship between grey matter volume and intelligence only develops over time, as no significant positive relationship can be found between grey matter volume and intelligence in children under 11.

An underlying caveat to research into the relationship of grey matter volume and intelligence is demonstrated by the hypothesis of neural efficiency. The findings that more intelligent individuals are more efficient at using their neurons might indicate that the correlation of grey matter to intelligence reflects selective elimination of unused synapses, and thus a better brain circuitry.

White matter

Similar to grey matter, white matter has been shown to correlate positively with intelligence in humans. White matter consists mainly of myelinated neuronal axons, responsible for delivering signals between neurons. The pinkish-white color of white matter is actually a result of these myelin sheaths that electrically insulate neurons that are transmitting signals to other neurons. White matter connects different regions of grey matter in the cerebrum together. These interconnections make transport more seamless and allow us to perform tasks easier. Significant correlations between intelligence and the corpus callosum have been found, as larger callosal areas have been positively correlated with cognitive performance. However, there appear to be differences in importance for white matter between verbal and nonverbal intelligence, as although both verbal and nonverbal measures of intelligence correlate positively with the size of the corpus callosum, the correlation for intelligence and corpus callosum size was larger (.47) for nonverbal measures than that for verbal measures (.18). Anatomical mesh-based geometrical modelling has also shown positive correlations between the thickness of the corpus callosum and Intelligence in healthy adults.

White matter integrity has also been found to be related to Intelligence. White matter tract integrity is important for information processing speed, and therefore reduced white matter integrity is related to lower intelligence. The effect of white matter integrity is mediate entirely through information processing speed. These findings indicate that the brain is structurally interconnected and that axonal fibres are integrally important for fast information process, and thus general intelligence.

Contradicting the findings described above, VBM failed to find a relationship between the corpus callosum and intelligence in healthy adults. This contradiction can be viewed to signify that the relationship between white matter volume and intelligence is not as robust as that of grey matter and intelligence.

Cortical thickness

Cortical thickness has also been found to correlate positively with intelligence in humans. However, the rate of growth of cortical thickness is also related to intelligence. In early childhood, cortical thickness displays a negative correlation with intelligence, while by late childhood this correlation has shifted to a positive one. More intelligent children were found to develop cortical thickness more steadily and over longer periods of time than less bright children. Studies have found cortical thickness to explain 5% in the variance of intelligence among individuals. In a study conducted to find associations between cortical thickness and general intelligence between different groups of people, sex did not play a role in intelligence. Although it is hard to pin intelligence on age based on cortical thickness due to different socioeconomic circumstances and education levels, older subjects (17 - 24) tended to have less variances in terms of intelligence than when compared to younger subjects (19 - 17).

Cortical convolution

Cortical convolution has increased the folding of the brain’s surface over the course of human evolution. It has been hypothesized that the high degree of cortical convolution may be a neurological substrate that supports some of the human brain's most distinctive cognitive abilities. Consequently, individual intelligence within the human species might be modulated by the degree of cortical convolution.

An analysis published in 2019 found the contours of 677 kids' brain had a genetic correlation of almost 1 between IQ and surface area of the supramarginal gyrus on the left side of the brain.

Neural efficiency

The neural efficiency hypothesis postulates that more intelligent individuals display less activation in the brain during cognitive tasks, as measured by Glucose metabolism. A small sample of participants (N=8) displayed negative correlations between intelligence and absolute regional metabolic rates ranging from -0.48 to -0.84, as measured by PET scans, indicating that brighter individuals were more effective processors of information, as they use less energy. According to an extensive review by Neubauer & Fink a large number of studies (N=27) have confirmed this finding using methods such as PET scans, EEG and fMRI.

fMRI and EEG studies have revealed that task difficulty is an important factor affecting neural efficiency. More intelligent individuals display neural efficiency only when faced with tasks of subjectively easy to moderate difficulty, while no neural efficiency can be found during difficult tasks. In fact, more able individuals appear to invest more cortical resources in tasks of high difficulty. This appears to be especially true for the Prefrontal Cortex, as individuals with higher intelligence displayed increased activation of this area during difficult tasks compared to individuals with lower intelligence. It has been proposed that the main reason for the neural efficiency phenomenon could be that individuals with high intelligence are better at blocking out interfering information than individuals with low intelligence.

Further research

Some scientists prefer to look at more qualitative variables to relate to the size of measurable regions of known function, for example relating the size of the primary visual cortex to its corresponding functions, that of visual performance.

In a study of the head growth of 633 term-born children from the Avon Longitudinal Study of Parents and Children cohort, it was shown that prenatal growth and growth during infancy were associated with subsequent IQ. The study’s conclusion was that the brain volume a child achieves by the age of 1 year helps determine later intelligence. Growth in brain volume after infancy may not compensate for poorer earlier growth.

There is an association between IQ and myopia. One suggested explanation is that one or several pleiotropic gene(s) affect the size of the neocortex part of the brain and eyes simultaneously.

Parieto-frontal integration theory

In 2007, Behavioral and Brain Sciences published a target article that put forth a biological model of intelligence based on 37 peer-reviewed neuroimaging studies (Jung & Haier, 2007). Their review of a wealth of data from functional imaging (functional magnetic resonance imaging and positron emission tomography) and structural imaging (diffusion MRI, voxel-based morphometry, in vivo magnetic resonance spectroscopy) argues that human intelligence arises from a distributed and integrated neural network comprising brain regions in the frontal and parietal lobes.

A recent lesion mapping study conducted by Barbey and colleagues provides evidence to support the P-FIT theory of intelligence.

Brain injuries at an early age isolated to one side of the brain typically results in relatively spared intellectual function and with IQ in the normal range.

Primates

Brain size

Another theory of brain size in vertebrates is that it may relate to social rather than mechanical skill. Cortical size relates directly to a pairbonding life style and among primates cerebral cortex size varies directly with the demands of living in a large complex social network. Compared to other mammals, primates have significantly larger brain size. Additionally, most primates are found to be polygynandrous, having many social relationships with others. Although inconclusive, some studies have shown that this polygnandrous statue correlates to brain size.

Intelligence in chimpanzees has been found to be relate to brain size, grey matter volume, and cortical thickness, as in humans.

Health

Several environmental factors related to health can lead to significant cognitive impairment, particularly if they occur during pregnancy and childhood when the brain is growing and the blood–brain barrier is less effective. Developed nations have implemented several health policies regarding nutrients and toxins known to influence cognitive function. These include laws requiring fortification of certain food products and laws establishing safe levels of pollutants (e.g. lead, mercury, and organochlorides). Comprehensive policy recommendations targeting reduction of cognitive impairment in children have been proposed.

Evolution of human intelligence

From Wikipedia, the free encyclopedia
 
The evolution of human intelligence is closely tied to the evolution of the human brain and to the origin of language. The timeline of human evolution spans approximately 7 million years, from the separation of the genus Pan until the emergence of behavioral modernity by 50,000 years ago. The first 3 million years of this timeline concern Sahelanthropus, the following 2 million concern Australopithecus and the final 2 million span the history of the genus Homo in the Paleolithic era.

Many traits of human intelligence, such as empathy, theory of mind, mourning, ritual, and the use of symbols and tools, are apparent in great apes although in less sophisticated forms than found in humans, such as great ape language.

History

Hominidae

The great apes (hominidae) show considerable cognitive and empathic abilities. Chimpanzees can make tools and use them to acquire foods and for social displays; they have sophisticated hunting strategies requiring cooperation, influence and rank; they are status conscious, manipulative and capable of deception; they can learn to use symbols and understand aspects of human language including some relational syntax, concepts of number and numerical sequence.

Homininae

Chimpanzee mother and baby
 
Around 10 million years ago, the Earth's climate entered a cooler and drier phase, which led eventually to the Quaternary glaciation beginning some 2.6 million years ago. One consequence of this was that the north African tropical forest began to retreat, being replaced first by open grasslands and eventually by desert (the modern Sahara). As their environment changed from continuous forest to patches of forest separated by expanses of grassland, some primates adapted to a partly or fully ground-dwelling life. Here they were exposed to predators, such as the big cats, from whom they had previously been safe. 

These environmental pressures caused selection to favor bipedalism: walking on hind legs. This gave the Homininae's eyes greater elevation, the ability to see approaching danger further off, and a more efficient means of locomotion. It also freed the arms from the task of walking and made the hands available for tasks such as gathering food. At some point the bipedal primates developed handedness, giving them the ability to pick up sticks, bones and stones and use them as weapons, or as tools for tasks such as killing smaller animals, cracking nuts, or cutting up carcasses. In other words, these primates developed the use of primitive technology. Bipedal tool-using primates form the Hominina subtribe, of which the earliest species, such as Sahelanthropus tchadensis, date to about 7 to 5 million years ago. 

From about 5 million years ago, the hominin brain began to develop rapidly in both size and differentiation of function. There has been a gradual increase in brain volume as humans progressed along the timeline of evolution, starting from about 600 cm3 in Homo habilis up to 1500 cm3 in Homo neanderthalensis. Thus, in general there's a correlation between brain volume and intelligence. However, modern Homo sapiens have a brain volume slightly smaller (1250 cm3) than neanderthals, and the Flores hominids (Homo floresiensis), nicknamed hobbits, had a cranial capacity of about 380 cm3 (considered small for a chimpanzee) about a third of that of H. erectus. It is proposed that they evolved from H. erectus as a case of insular dwarfism. With their three times smaller brain the Flores hominids apparently used fire and made tools as sophisticated as those of their ancestor H.erectus. In this case, it seems that for intelligence, the structure of the brain is more important than its volume.

Homo

Roughly 2.4 million years ago Homo habilis had appeared in East Africa: the first known human species, and the first known to make stone tools, yet the disputed findings of signs of tool use from even earlier ages and from the vicinity as multiple Australopithecus fossils may put this to question its "greater intelligence when compared to earlier and more primitive Australopithecus genus". 

The use of tools conferred a crucial evolutionary advantage, and required a larger and more sophisticated brain to co-ordinate the fine hand movements required for this task. Our knowledge of the complexity of behaviour of Homo habilis is not limited to stone culture, they also had habitual therapic use of toothpicks. The evolution of a larger brain created a problem for early humans, however. A larger brain requires a larger skull, and thus requires the female to have a wider birth canal for the newborn's larger skull to pass through. But if the female's birth canal grew too wide, her pelvis would be so wide that she would lose the ability to run, which was a necessary skill 2 million years ago.

The solution to this was to give birth at an early stage of fetal development, before the skull grew too large to pass through the birth canal. This adaptation enabled the human brain to continue to grow, but it imposed a new discipline. The need to care for helpless infants for long periods of time forced humans to become less mobile. Human bands increasingly stayed in one place for long periods, so that females could care for infants, while males hunted food and fought with other bands that competed for food sources. As a result, humans became even more dependent on tool-making to compete with other animals and other humans, and relied less on body size and strength.

About 200,000 years ago Europe and the Middle East were colonized by Neanderthal man, extinct by 39,000 years ago following the appearance of modern humans in the region from 40,000–45,000 years ago.

Homo sapiens

"The Lion-man", found in the Hohlenstein-Stadel cave of Germany's Swabian Alb and dated to 40,000 years ago, is associated with the Aurignacian culture and is the oldest known anthropomorphic animal figurine in the world.
 
Quaternary extinction eventQuaternary extinction eventHolocene extinctionHolocene extinctionYellowstone CalderaYellowstone CalderaToba catastrophe theoryHomo heidelbergensisHomo neanderthalensisHomo antecessorHomo sapiensHomo habilisHomo georgicusHomo ergasterHomo erectusHomoHomo
Dates approximate, consult articles for details (From 2000000 BC till 2013 AD in (partial) exponential notation)

Homo sapiens intelligence

Around 200,000 years ago, Homo sapiens first appeared in East Africa. It is unclear to what extent these early modern humans had developed language, music, religion etc. They spread throughout Africa over the following approximately 50,000 years.

According to proponents of the Toba catastrophe theory, the climate in non-tropical regions of the earth experienced a sudden freezing about 70,000 years ago, because of a huge explosion of the Toba volcano that filled the atmosphere with volcanic ash for several years. This reduced the human population to less than 10,000 breeding pairs in equatorial Africa, from which all modern humans are descended. Being unprepared for the sudden change in climate, the survivors were those intelligent enough to invent new tools and ways of keeping warm and finding new sources of food (for example, adapting to ocean fishing based on prior fishing skills used in lakes and streams that became frozen).

Around 80,000–100,000 years ago, three main lines of Homo sapiens diverged, bearers of mitochondrial haplogroup L1 (mtDNA) / A (Y-DNA) colonizing Southern Africa (the ancestors of the Khoisan/Capoid peoples), bearers of haplogroup L2 (mtDNA) / B (Y-DNA) settling Central and West Africa (the ancestors of Niger–Congo and Nilo-Saharan speaking peoples), while the bearers of haplogroup L3 remained in East Africa.

The "Great Leap Forward" leading to full behavioral modernity sets in only after this separation. Rapidly increasing sophistication in tool-making and behaviour is apparent from about 80,000 years ago, and the migration out of Africa follows towards the very end of the Middle Paleolithic, some 60,000 years ago. Fully modern behaviour, including figurative art, music, self-ornamentation, trade, burial rites etc. is evident by 30,000 years ago. The oldest unequivocal examples of prehistoric art date to this period, the Aurignacian and the Gravettian periods of prehistoric Europe, such as the Venus figurines and cave painting (Chauvet Cave) and the earliest musical instruments (the bone pipe of Geissenklösterle, Germany, dated to about 36,000 years ago).

The human brain has evolved gradually over the passage of time; a series of incremental changes occurred as a result of external stimuli and conditions. It is crucial to keep in mind that evolution operates within a limited framework at a given point in time. In other words, the adaptations that a species can develop are not infinite and are defined by what has already taken place in the evolutionary timeline of a species. Given the immense anatomical and structural complexity of the brain, its evolution (and the congruent evolution of human intelligence), can only be reorganized in a finite number of ways. The majority of said changes occur either in terms of size or in terms of developmental timeframes.

Motor and sensory areas of the cerebral cortex; dashed areas shown are commonly left hemisphere dominant.
 
There have been studies that strongly support the idea that the level of intelligence associated with humans is not unique to our species. Scholars suggest that this could have, in part, been caused by convergent evolution. One common characteristic that is present in species of "high degree intelligence" (i.e. dolphins, great apes, and humans - Homo sapiens) is a brain of enlarged size. Along with this, there is a more developed neocortex, a folding of the cerebral cortex, and von Economo neurons. Said neurons are linked to social intelligence and the ability to gauge what another is thinking or feeling and, interestingly, are also present in bottlenose dolphins. The cerebral cortex is divided into four lobes (frontal, parietal, occipital, and temporal) each with specific functions. The cerebral cortex is significantly larger in humans than in any other animal and is responsible for higher thought processes such as: reasoning, abstract thinking, and decision making.

Another characteristic that sets humans apart from any other species is the ability to produce and understand complex, syntactic language. The cerebral cortex, particularly in the temporal, parietal, and frontal lobes, are populated with neural circuits dedicated to language. There are two main areas of the brain commonly associated with language, namely: Wernicke's area and Broca's area. The former is responsible for the understanding of speech and the latter for the production of speech. Homologous regions have been found in other species (i.e. Area 44 and 45 have been studied in chimpanzees) but they are not as strongly related to or involved in linguistic activities as in humans.

A big portion of the scholarly literature focus on the evolution, and subsequent influence, of culture. This is in part because the leaps human intelligence has taken are far greater than those that would have resulted if our ancestors had simply responded to their environments, inhabiting them as hunter-gatherers. (Richardson 273). 

In short, the complexity and marvel of human intelligence only emerge inside of a specific culture and history. Selection for cooperation aided our ancestors in surviving harsh ecological conditions and did so by creating a specific type of intelligence. An intelligence that, today, is highly variant from individual to individual.

Models

Social brain hypothesis

The social brain hypothesis was proposed by British anthropologist Robin Dunbar, who argues that human intelligence did not evolve primarily as a means to solve ecological problems, but rather as a means of surviving and reproducing in large and complex social groups. Some of the behaviors associated with living in large groups include reciprocal altruism, deception and coalition formation. These group dynamics relate to Theory of Mind or the ability to understand the thoughts and emotions of others, though Dunbar himself admits in the same book that it is not the flocking itself that causes intelligence to evolve (as shown by ruminants).

Dunbar argues that when the size of a social group increases, the number of different relationships in the group may increase by orders of magnitude. Chimpanzees live in groups of about 50 individuals whereas humans typically have a social circle of about 150 people, which is also the typical size of social communities in small societies and personal social networks; this number is now referred to as Dunbar's number. In addition, there is evidence to suggest that the success of groups is dependent on their size at foundation, with groupings of around 150 being particularly successful, potentially reflecting the fact that communities of this size strike a balance between the minimum size of effective functionality and the maximum size for creating a sense of commitment to the community. According to the social brain hypothesis, when hominids started living in large groups, selection favored greater intelligence. As evidence, Dunbar cites a relationship between neocortex size and group size of various mammals.

Criticism

Phylogenetic studies of brain sizes in primates show that while diet predicts primate brain size, sociality does not predict brain size when corrections are made for cases in which diet affects both brain size and sociality. The exceptions to the predictions of the social intelligence hypothesis, which that hypothesis has no predictive model for, are successfully predicted by diets that are either nutritious but scarce or abundant but poor in nutrients. Researchers have found that frugivores tend to exhibit larger brain size than folivores. One potential explanation for this finding is that frugivory requires 'extractive foraging,' or the process of locating and preparing hard-shelled foods, such as nuts, insects, and fruit. Extractive foraging requires higher cognitive processing, which could help explain larger brain size. However, other researchers argue that extractive foraging was not a catalyst in the evolution of primate brain size, demonstrating that some non primates exhibit advanced foraging techniques. Other explanations for the positive correlation between brain size and frugivory highlight how the high-energy, frugivore diet facilitates fetal brain growth and requires spatial mapping to locate the embedded foods.

Meerkats have far more social relationships than their small brain capacity would suggest. Another hypothesis is that it is actually intelligence that causes social relationships to become more complex, because intelligent individuals are more difficult to learn to know.

There are also studies that show that Dunbar's number is not the upper limit of the number of social relationships in humans either.

The hypothesis that it is brain capacity that sets the upper limit for the number of social relationships is also contradicted by computer simulations that show simple unintelligent reactions to be sufficient to emulate "ape politics" and by the fact that some social insects such as the paper wasp do have hierarchies in which each individual has its place (as opposed to herding without social structure) and maintains their hierarchies in groups of approximately 80 individuals with their brains smaller than that of any mammal.

Reduction in aggression

Another theory that tries to explain the growth of human intelligence is the reduced aggression theory (aka self-domestication theory). According to this strand of thought what led to the evolution of advanced intelligence in Homo sapiens was a drastic reduction of the aggressive drive. This change separated us from other species of monkeys and primates, where this aggressivity is still in plain sight, and eventually lead to the development of quintessential human traits such as empathy, social cognition and culture. This theory has received strong support from studies of animal domestication where selective breeding for tameness has, in only a few generations, led to the emergence of impressive "humanlike" abilities. Tamed foxes, for example, exhibit advanced forms of social communication (following pointing gestures), pedomorphic physical features (childlike faces, floppy ears) and even rudimentary forms of theory of mind (eye contact seeking, gaze following). Evidence also comes from the field of ethology (which is the study of animal behavior, focused on observing species in their natural habitat rather than in controlled laboratory settings) where it has been found that animals with a gentle and relaxed manner of interacting with each other – like for example stumptailed macaques, orangutans and bonobos – have more advanced socio-cognitive abilities than those found among the more aggressive chimpanzees and baboons. It is hypothesized that these abilities derive from a selection against aggression.

On a mechanistic level these changes are believed to be the result of a systemic downregulation of the sympathetic nervous system (the fight-or-flight reflex). Hence, tamed foxes show a reduced adrenal gland size and have an up to fivefold reduction in both basal and stress-induced blood cortisol levels. Similarly, domesticated rats and guinea pigs have both reduced adrenal gland size and reduced blood corticosterone levels. It seems as though the neoteny of domesticated animals significantly prolongs the immaturity of their hypothalamic-pituitary-adrenal system (which is otherwise only immature for a short period when they are pups/kittens) and this opens up a larger "socialization window" during which they can learn to interact with their caretakers in a more relaxed way. 

This downregulation of sympathetic nervous system reactivity is also believed to be accompanied by a compensatory increase in a number of opposing organs and systems. Although these are not as well specified various candidates for such "organs" have been proposed: the parasympathetic system as a whole, the septal area over the amygdala, the oxytocin system, the endogenous opioids and various forms of quiescent immobilization which antagonize the fight-or-flight reflex.

Social exchange theory

Other studies suggest that social exchange between individuals is a vital adaptation to the human brain, going as far to say that the human mind could be equipped with a neurocognitive system specialized for reasoning about social change. Social Exchange is a vital adaptation that evolved in social species and has become exceptionally specialized in humans.This adaption will develop by natural selection when two parties can make themselves better off than they were before by exchanging things one party values less for things the other party values for more. However, selection will only pressure social exchange when both parties are receiving mutual benefits from their relative situation; if one party cheats the other by receiving a benefit while the other is harmed, then selection will stop. Consequently, the existence of cheaters—those who fail to deliver fair benefits—threatens the evolution of exchange. Using evolutionary game theory, it has been shown that adaptations for social exchange can be favored and stably maintained by natural selection, but only if they include design features that enable them to detect cheaters, and cause them to channel future exchanges to reciprocators and away from cheaters. Thus, humans use social contracts to lay the benefits and losses each party will be receiving (if you accept benefit B from me, then you must satisfy my requirement R). Humans have evolved an advanced cheater detection system, equipped with proprietary problem-solving strategies that evolved to match the recurrent features of their corresponding problem domains. Not only do humans need to determine that the contract was violated, but also if the violation was intentionally done. Therefore, systems are specialized to detect contract violations that imply intentional cheating.

One problem with the hypothesis that specific punishment for intentional deception could coevolve with intelligence is the fact that selective punishment of individuals with certain characteristics selects against the characteristics in question. For example, if only individuals capable of remembering what they had agreed to were punished for breaking agreements, evolution would have selected against the ability to remember what one had agreed to. Though this becomes a superficial argument after considering the balancing positive selection for the ability to successfully 'make ones case'. Intelligence predicts the number of arguments one can make when taking either side of a debate. Humans who could get away with behaviours that exploited within and without-group cooperation, getting more while giving less, would overcome this.

Sexual selection

This model, which invokes sexual selection, is proposed by Geoffrey Miller who argues that human intelligence is unnecessarily sophisticated for the needs of hunter-gatherers to survive. He argues that the manifestations of intelligence such as language, music and art did not evolve because of their utilitarian value to the survival of ancient hominids. Rather, intelligence may have been a fitness indicator. Hominids would have been chosen for greater intelligence as an indicator of healthy genes and a Fisherian runaway positive feedback loop of sexual selection would have led to the evolution of human intelligence in a relatively short period.

In many species, only males have impressive secondary sexual characteristics such as ornaments and show-off behavior, but sexual selection is also thought to be able to act on females as well in at least partially monogamous species. With complete monogamy, there is assortative mating for sexually selected traits. This means that less attractive individuals will find other less attractive individuals to mate with. If attractive traits are good fitness indicators, this means that sexual selection increases the genetic load of the offspring of unattractive individuals. Without sexual selection, an unattractive individual might find a superior mate with few deleterious mutations, and have healthy children that are likely to survive. With sexual selection, an unattractive individual is more likely to have access only to an inferior mate who is likely to pass on many deleterious mutations to their joint offspring, who are then less likely to survive.

Sexual selection is often thought to be a likely explanation for other female-specific human traits, for example breasts and buttocks far larger in proportion to total body size than those found in related species of ape. It is often assumed that if breasts and buttocks of such large size were necessary for functions such as suckling infants, they would be found in other species. That human female breasts (typical mammalian breast tissue is small) are found sexually attractive by many men is in agreement with sexual selection acting on human females secondary sexual characteristics. 

Sexual selection for intelligence and judging ability can act on indicators of success, such as highly visible displays of wealth. Growing human brains require more nutrition than brains of related species of ape. It is possible that for females to successfully judge male intelligence, they must be intelligent themselves. This could explain why despite the absence of clear differences in intelligence between males and females on average, there are clear differences between male and female propensities to display their intelligence in ostentatious forms.

This absence of difference is now known to exist at the middle of distributions. Average intelligence doesn't differ much between genders, but because female selection is restricted more towards males at the top end of male-male hierarchies or those increasingly above average in physical attractiveness, male trait distributions often have longer tails; that is to say the lowest and highest intelligences (and many more traits) in male populations extend further out into the lowest and highest values of the distribution than for female traits. This is because it paid to be a highly variable male, as average males would have consistently low opportunity, but variable males had a chance of falling on the preferred side of the trait distribution.

Critique

The sexual selection by the disability principle/fitness display model of the evolution of human intelligence is criticized by certain researchers for issues of timing of the costs relative to reproductive age. While sexually selected ornaments such as peacock feathers and moose antlers develop either during or after puberty, timing their costs to a sexually mature age, human brains expend large amounts of nutrients building myelin and other brain mechanisms for efficient communication between the neurons early in life. These costs early in life build facilitators that reduce the cost of neuron firing later in life, and as a result the peaks of the brain's costs and the peak of the brain's performance are timed on opposite sides of puberty with the costs peaking at a sexually immature age while performance peaks at a sexually mature age. Critical researchers argue that this means that the costs that intelligence is a signal of reduce the chances of surviving to reproductive age, does not signal fitness of sexually mature individuals and, since the disability principle is about selection for disabilities in sexually immature individuals that evolutionarily increase the offspring's chance of surviving to reproductive age, would be selected against and not for by its mechanisms. These critics argue that human intelligence evolved by natural selection citing that unlike sexual selection, natural selection have produced many traits that cost the most nutrients before puberty including immune systems and accumulation and modification for increased toxicity of poisons in the body as a protective measure against predators.

Intelligence as a disease-resistance sign

The number of people with severe cognitive impairment caused by childhood viral infections like meningitis, protists like Toxoplasma and Plasmodium, and animal parasites like intestinal worms and schistosomes is estimated to be in the hundreds of millions. Even more people live with moderate mental damages, such as inability to complete difficult tasks, that are not classified as 'diseases' by medical standards, may still be considered as inferior mates by potential sexual partners.

Thus, widespread, virulent, and archaic infections are greatly involved in natural selection for cognitive abilities. People infected with parasites may have brain damage and obvious maladaptive behavior in addition to visible signs of disease. Smarter people can more skillfully learn to distinguish safe non-polluted water and food from unsafe kinds and learn to distinguish mosquito infested areas from safe areas. Smarter people can more skillfully find and develop safe food sources and living environments. Given this situation, preference for smarter child-bearing/rearing partners increases the chance that their descendants will inherit the best resistance alleles, not only for immune system resistance to disease, but also smarter brains for learning skills in avoiding disease and selecting nutritious food. When people search for mates based on their success, wealth, reputation, disease-free body appearance, or psychological traits such as benevolence or confidence; the effect is to select for superior intelligence that results in superior disease resistance.

Ecological dominance-social competition model

A predominant model describing the evolution of human intelligence is ecological dominance-social competition (EDSC), explained by Mark V. Flinn, David C. Geary and Carol V. Ward based mainly on work by Richard D. Alexander. According to the model, human intelligence was able to evolve to significant levels because of the combination of increasing domination over habitat and increasing importance of social interactions. As a result, the primary selective pressure for increasing human intelligence shifted from learning to master the natural world to competition for dominance among members or groups of its own species.

As advancement, survival and reproduction within an increasing complex social structure favored ever more advanced social skills, communication of concepts through increasingly complex language patterns ensued. Since competition had shifted bit by bit from controlling "nature" to influencing other humans, it became of relevance to outmaneuver other members of the group seeking leadership or acceptance, by means of more advanced social skills. A more social and communicative person would be more easily selected.

Intelligence dependent on brain size

Human intelligence is developed to an extreme level that is not necessarily adaptive in an evolutionary sense. Firstly, larger-headed babies are more difficult to give birth to and large brains are costly in terms of nutrient and oxygen requirements. Thus the direct adaptive benefit of human intelligence is questionable at least in modern societies, while it is difficult to study in prehistoric societies. Since 2005, scientists have been evaluating genomic data on gene variants thought to influence head size, and have found no evidence that those genes are under strong selective pressure in current human populations. The trait of head size has become generally fixed in modern human beings.

While decreased brain size has strong correlation with lower intelligence in humans, some modern humans have brain sizes as small as Homo Erectus but normal intelligence (based on IQ tests) for modern humans. Increased brain size in humans may allow for greater capacity for specialized expertise.

Expanded cortical regions

The two major perspectives on primate brain evolution are the concerted and mosaic approaches. In the concerted evolution approach, cortical expansions in the brain are considered to be a by-product of a larger brain, rather than adaptive potential. Studies have supported the concerted evolution model by finding cortical expansions between macaques and marmosets are comparable to that of humans and macaques. Researchers attribute this result to the constraints on the evolutionary process of increasing brain size. In the mosaic approach, cortical expansions are attributed to their adaptive advantage for the species. Researchers have attributed hominin evolution to mosaic evolution.

Simian primate brain evolution studies show that specific cortical regions associated with high-level cognition have demonstrated the greatest expansion over primate brain evolution. Sensory and motor regions have showcased limited growth. Three regions associated with complex cognition include the frontal lobe, temporal lobe, and the medial wall of the cortex. Studies demonstrate that the enlargement in these regions is disproportionately centered in the temporoparietal junction (TPJ), lateral prefrontal cortex (LPFC), and anterior cingulate cortex (ACC). The TPJ is located in the parietal lobe and is associated with morality, theory of mind, and spatial awareness. Additionally, the Wernicke's area is located in the TPJ. Studies have suggested that the region assists in language production, as well as language processing. The LPFC is commonly associated with planning and working memory functions. The Broca's area, the second major region associated with language processing, is also located in the LPFC. The ACC is associated with detecting errors, monitoring conflict, motor control, and emotion. Specifically, researchers have found that the ACC in humans is disproportionately expanded when compared to the ACC in macaques.

Studies on cortical expansions in the brain have been used to examine the evolutionary basis of neurological disorders, such as Alzheimer's disease. For example, researchers associate the expanded TPJ region with Alzheimer's disease. However, other researchers found no correlation between expanded cortical regions in the human brain and the development of Alzheimer's disease.
Cellular, genetic, and circuitry changes
Human brain evolution involves cellular, genetic, and circuitry changes. On a genetic level, humans have a modified FOXP2 gene, which is associated with speech and language development. The human variant of the gene SRGAP2, SRGAP2C, enables greater dendritic spine density which fosters greater neural connections. On a cellular level, studies demonstrate von Economo neurons (VENs) are more prevalent in humans than other primates. Studies show that VENs are associated with empathy, social awareness and self-control. Studies show that the striatum plays a role in understanding reward and pair-bond formation. On a circuitry level, humans exhibit a more complex mirror neuron system, greater connection between the two major language processing areas (Wernicke's area and Broca's area), and a vocal control circuit that connects the motor cortex and brain stem. The mirror neuron system is associated with social cognition, theory of mind, and empathy. Studies have demonstrated the presence of the mirror neuron system in both macaques in humans; However, the mirror neuron system is only activated in macaques when observing transitive movements.

Group selection

Group selection theory contends that organism characteristics that provide benefits to a group (clan, tribe, or larger population) can evolve despite individual disadvantages such as those cited above. The group benefits of intelligence (including language, the ability to communicate between individuals, the ability to teach others, and other cooperative aspects) have apparent utility in increasing the survival potential of a group. 

In addition, the theory of group selection is inherently tied to Darwin's theory of natural selection. Specifically, that "group-related adaptations must be attributed to the natural selection of alternative groups of individuals and that the natural selection of alternative alleles within populations will be opposed to this development".

Between-group selection can be used to explain the changes and adaptations that arise within a group of individuals. Group-related adaptations and changes are a byproduct of between-group selection as traits or characteristics that prove to be advantageous in relation to another group will become increasingly popular and disseminated within a group. In the end, increasing its overall chance of surviving a competing group. 

However, this explanation cannot be applied to humans (and other species, predominantly other mammals) that live in stable, established social groupings. This is because of the social intelligence that functioning within these groups requires from the individual. Humans, while they are not the only ones, possess the cognitive and mental capacity to form systems of personal relationships and ties that extend well beyond those of the nucleus of family. The continuous process of creating, interacting, and adjusting to other individuals is a key component of many species' ecology. 

These concepts can be tied to the social brain hypothesis, mentioned above. This hypothesis posits that human cognitive complexity arose as a result of the higher level of social complexity required from living in enlarged groups. These bigger groups entail a greater amount of social relations and interactions thus leading to a expanded quantity of intelligence in humans. However, this hypothesis has been under academic scrutiny in recent years and has been largely disproven. In fact, the size of a species' brain can be much better predicted by diet instead of measures of sociality as noted by the study conducted by DeCasien et. al. They found that ecological factors (such as: folivory/frugivory, environment) explain a primate brain size much better than social factors (such as: group size, mating system).

Nutritional status

Diets deficient in iron, zinc, protein, iodine, B vitamins, omega 3 fatty acids, magnesium and other nutrients can result in lower intelligence either in the mother during pregnancy or in the child during development. While these inputs did not have an effect on the evolution of intelligence they do govern its expression. A higher intelligence could be a signal that an individual comes from and lives in a physical and social environment where nutrition levels are high, whereas a lower intelligence could imply a child, its mother, or both, come from a physical and social environment where nutritional levels are low. Previc emphasizes the contribution of nutritional factors, especially meat and shellfish consumption, to elevations of dopaminergic activity in the brain, which may have been responsible for the evolution of human intelligence since dopamine is crucial to working memory, cognitive shifting, abstract, distant concepts, and other hallmarks of advanced intelligence.

Operator (computer programming)

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