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Friday, October 26, 2018

Developmental psychology

From Wikipedia, the free encyclopedia

Special used in the psychological study of infants
 
Piaget's test for Conservation. One of the many experiments used for children
 
Developmental psychology is the scientific study of how and why human beings change over the course of their life. Originally concerned with infants and children, the field has expanded to include adolescence, adult development, aging, and the entire lifespan. Developmental psychologists aim to explain how thinking, feeling, and behaviors change throughout life. This field examines change across three major dimensions: physical development, cognitive development, and socioemotional development. Within these three dimensions are a broad range of topics including motor skills, executive functions, moral understanding, language acquisition, social change, personality, emotional development, self-concept, and identity formation.

Developmental psychology examines the influences of nature and nurture on the process of human development, and processes of change in context and across time. Many researchers are interested in the interactions among personal characteristics, the individual's behavior and environmental factors, including the social context and the built environment. Ongoing debates include biological essentialism vs. neuroplasticity and stages of development vs. dynamic systems of development.

Developmental psychology involves a range of fields, such as educational psychology, child psychopathology, forensic developmental psychology, child development, cognitive psychology, ecological psychology, and cultural psychology. Influential developmental psychologists from the 20th century include Urie Bronfenbrenner, Erik Erikson, Sigmund Freud, Jean Piaget, Barbara Rogoff, Esther Thelen, and Lev Vygotsky.

Historical antecedents

John B. Watson and Jean-Jacques Rousseau are typically cited as providing the foundations for modern developmental psychology. In the mid-18th century Jean Jacques Rousseau described three stages of development: infants (infancy), puer (childhood) and adolescence in Emile: Or, On Education. Rousseau's ideas were taken up strongly by educators at the time.

It generally focuses on how and why certain modifications throughout an individual’s life-cycle (cognitive, social, intellectual, personality) and human growth change over time. There are many theorists that have made a profound contribution to this area of psychology. For example, Erik Erikson developed a model of eight stages of psychological development. He believed that humans developed in stages throughout their lifetimes and this would affect their behaviors. (Similar ideas to Sigmund Freud.)

In the late 19th century, psychologists familiar with the evolutionary theory of Darwin began seeking an evolutionary description of psychological development; prominent here was the pioneering psychologist G. Stanley Hall, who attempted to correlate ages of childhood with previous ages of humanity. James Mark Baldwin who wrote essays on topics that included Imitation: A Chapter in the Natural History of Consciousness and Mental Development in the Child and the Race: Methods and Processes. Baldwin was heavily involved in the theory of developmental psychology. Sigmund Freud, whose concepts were developmental, significantly affected public perceptions.

Theories

Psychosexual development

Sigmund Freud believed that we all had a conscious, preconscious, and unconscious level. In the conscious, we are aware of our mental process. The preconscious involves information that, though not currently in our thoughts, can be brought into consciousness. Lastly, the unconscious includes mental processes we are unaware of.

He believed there is tension between the conscious and unconscious because the conscious tries to hold back what the unconscious tries to express. To explain this he developed three personality structures: the id, ego, and superego. The id, the most primitive of the three, functions according to the pleasure principle: seek pleasure and avoid pain. The superego plays the critical and moralizing role; and the ego is the organized, realistic part that mediates between the desires of the id and the superego.

Based on this, he proposed five universal stages of development, that each is characterized by the erogenous zone that is the source of the child's psychosexual energy. The first is the oral stage, which occurs from birth to 12 months of age. During the oral stage, "the libido is centered in a baby's mouth." The baby is able to suck. The second is the anal stage, from one to three years of age. During the anal stage, the child defecates from the anus and is often fascinated with their defecation. The third is the phallic stage, which occurs from three to five years of age (most of a person's personality forms by this age). During the phallic stage, the child is aware of their sexual organs. The fourth is the latency stage, which occurs from age five until puberty. During the latency stage, the child's sexual interests are repressed. Stage five is the genital stage, which takes place from puberty until adulthood. During the genital stage, puberty starts happening.

Stages of moral development

Piaget claimed that logic and morality develop through constructive stages. Expanding on Piaget's work, Lawrence Kohlberg determined that the process of moral development was principally concerned with justice, and that it continued throughout the individual's lifetime.

He suggested three levels of moral reasoning; pre-conventional moral reasoning, conventional moral reasoning, and post-conventional moral reasoning. The pre-conventional moral reasoning is typical of children and is characterized by reasoning that is based on rewards and punishments associated with different courses of action. Conventional moral reason occurs during late childhood and early adolescence and is characterized by reasoning based on rules and conventions of society. Lastly, post-conventional moral reasoning is a stage during which the individual sees society's rules and conventions as relative and subjective, rather than as authoritative.

Kohlberg used the Heinz Dilemma to apply to his stages of moral development. The Heinz Dilemma involves Heinz's wife dying from cancer and Heinz having the dilemma to save his wife by stealing a drug. Preconventional morality, conventional morality, and post-conventional morality applies to Heinz's situation.

Stages of psychosocial development

German-American psychologist Erik Erikson and his collaborator and wife, Joan Erikson, conceptualized eight stages of psychosocial development that they theorized healthy individuals pass through as they develop from infancy to adulthood. The first stage is called "Trust vs. Mistrust" takes place in infancy. The best virtue for the first stage is hope, in the infant learning who to trust and having hope for a supportive group of people to be there for him/her. The second stage is "Autonomy vs. Shame and Doubt" with the best virtue being will. This takes place in early childhood where the child learns to become more independent by discovering what they are capable of where if the child is overly controlled, they believe to feel inadequate on surviving by themselves, which can lead to low self-esteem and doubt. The third stage is "Initiative vs. Guilt". The basic virtue that would be gained is the purpose and takes place in the play age. This is the stage where the child will be curious and have many interactions with other kids. They will ask many questions as their curiosity grows. If too much guilt is present, the child may have a slower and harder time interacting with other children. The fourth stage is "Industry (competence) vs. Inferiority". The basic virtue for this stage is competency which happens at the school age. This stage is when the child will try to win the approval of others and fit in and understand the value of their accomplishments. The fifth stage is "Identity vs. Role Confusion". The basic virtue gained is fidelity which takes place in adolescence. This is where the child will start to find who he/she is as a person in society. What sex role he/she picks. The sixth stage is "Intimacy vs. Isolation", which happens in young adults and the virtue gained is love. This is where the person will start to share his/her life with someone else intimately and emotionally. In not doing so, it could lead to isolation. The seventh stage is "Generativity vs. Stagnation". This happens in adulthood and the virtue gained would be care. We become stable and start to give back by raising a family and becoming involved in the community. The eighth stage is "Ego Integrity vs. Despair". This happens during maturity and wisdom is gained. When one grows old and they contemplate and look back and see the success or failure of their life. This is also the stage where one can also have closure and accept death without fearing anything.

Theories of cognitive development

Jean Piaget, a Swiss theorist, posited that children learn by actively constructing knowledge through hands-on experience. He suggested that the adult's role in helping the child learn was to provide appropriate materials that the child can interact with and use to construct. He used Socratic questioning to get children to reflect on what they were doing, and he tried to get them to see contradictions in their explanations.

Piaget believed that intellectual development takes place through a series of stages, which he described in his theory on cognitive development. Each stage consists of steps the child must master before moving to the next step. He believed that these stages are not separate from one another, but rather that each stage builds on the previous one in a continuous learning process. He proposed four stages: sensorimotor, pre-operational, concrete operational, and formal operational. Though he did not believe these stages occurred at any given age, many studies have determined when these cognitive abilities should take place.

Stages based on the model of hierarchical complexity

Michael Commons enhanced and simplified of Inhelder and Piaget's developmental and offers a standard method of examining the universal pattern of development. The Model of Hierarchical Complexity (MHC) is not based on the assessment of domain-specific information, It divides the Order of Hierarchical Complexity of tasks to be addressed from the Stage performance on those tasks. A stage is the order hierarchical complexity of the tasks the participant's successfully addresses. He expanded Piaget's original eight stage (counting the half stages) to fifteen stages. The stages are : 0 Calculatory; 1 Sensory & Motor; 2 Circular sensory-motor; 3 Sensory-motor; 4 Nominal; 5 Sentential; 6 Preoperational; 7 Primary; 8 Concrete; 9 Abstract; 10 Formal; 11 Systematic; 12 Metasystematic; 13 Paradigmatic; 14 Cross-paradigmatic; 15 Meta-Cross-paradigmatic. The order of hierarchical complexity of tasks predicts how difficult the performance is with an R ranging from 0.9 to 0.98.

In the MHC, there are three main axioms for an order to meet in order for the higher order task to coordinate the next lower order task. Axioms are rules that are followed to determine how the MHC orders actions to form a hierarchy. These axioms are: a) defined in terms of tasks at the next lower order of hierarchical complexity task action; b) defined as the higher order task action that organizes two or more less complex actions; that is, the more complex action specifies the way in which the less complex actions combine; c) defined as the lower order task actions have to be carried out non-arbitrarily.

Ecological systems theory

Bronfenbrenner's ecological systems theory

Ecological systems theory, originally formulated by Urie Bronfenbrenner, specifies four types of nested environmental systems, with bi-directional influences within and between the systems. The four systems are microsystem, mesosystem, exosystem, and macrosystem. Each system contains roles, norms and rules that can powerfully shape development. The microsystem is the direct environment in our lives such as our home and school. Mesosystem is how relationships connect to the microsystem. Exosystem is a larger social system where the child plays no role. Macrosystem refers to the cultural values, customs and laws of society.

The microsystem is the immediate environment surrounding and influencing the individual (example: school or the home setting). The mesosystem is the combination of two microsystems and how they influence each other (example: sibling relationships at home vs. peer relationships at school). The exosystem is the interaction among two or more settings that are indirectly linked (example: a father's job requiring more overtime ends up influencing his daughter's performance in school because he can no longer help with her homework). The macrosystem is broader taking into account social economic status, culture, beliefs, customs and morals (example: a child from a wealthier family sees a peer from a less wealthy family as inferior for that reason). Lastly, the chronosystem refers to the chronological nature of life events and how they interact and change the individual and their circumstances through transition (example: a mother losing her own mother to illness and no longer having that support in her life).

Since its publication in 1979, Bronfenbrenner's major statement of this theory, The Ecology of Human Development has had widespread influence on the way psychologists and others approach the study of human beings and their environments. As a result of this conceptualization of development, these environments—from the family to economic and political structures—have come to be viewed as part of the life course from childhood through to adulthood.

Zone of proximal development

Lev Vygotsky was a Russian theorist from the Soviet era, who posited that children learn through hands-on experience and social interactions with members of his/her culture. Unlike Piaget, he claimed that timely and sensitive intervention by adults when a child is on the edge of learning a new task (called the "zone of proximal development") could help children learn new tasks. This adult role is often referred to as the skilled "master," whereas the child is considered the learning apprentice through an educational process often termed "cognitive apprenticeship" Martin Hill stated that "The world of reality does not apply to the mind of a child." This technique is called "scaffolding," because it builds upon knowledge children already have with new knowledge that adults can help the child learn. Vygotsky was strongly focused on the role of culture in determining the child's pattern of development, arguing that development moves from the social level to the individual level. In other words, Vygotsky claimed that psychology should focus on the progress of human consciousness through the relationship of an individual and their environment. He felt that if scholars continued to disregard this connection, then this disregard would inhibit the full comprehension of the human consciousness.

Constructivism

Constructivism is a paradigm in psychology that characterizes learning as a process of actively constructing knowledge. Individuals create meaning for themselves or make sense of new information by selecting, organizing, and integrating information with other knowledge, often in the context of social interactions. Constructivism can occur in two ways: individual and social. Individual constructivism is when a person constructs knowledge through cognitive processes of their own experiences rather than by memorizing facts provided by others. Social constructivism is when individuals construct knowledge through an interaction between the knowledge they bring to a situation and social or cultural exchanges within that content.

Jean Piaget, a Swiss developmental psychologist, proposed that learning is an active process because children learn through experience and make mistakes and solve problems. Piaget proposed that learning should be whole by helping students understand that meaning is constructed.

Evolutionary developmental psychology

Evolutionary developmental psychology is a research paradigm that applies the basic principles of Darwinian evolution, particularly natural selection, to understand the development of human behavior and cognition. It involves the study of both the genetic and environmental mechanisms that underlie the development of social and cognitive competencies, as well as the epigenetic (gene-environment interactions) processes that adapt these competencies to local conditions.

EDP considers both the reliably developing, species-typical features of ontogeny (developmental adaptations), as well as individual differences in behavior, from an evolutionary perspective. While evolutionary views tend to regard most individual differences as the result of either random genetic noise (evolutionary byproducts) and/or idiosyncrasies (for example, peer groups, education, neighborhoods, and chance encounters) rather than products of natural selection, EDP asserts that natural selection can favor the emergence of individual differences via "adaptive developmental plasticity." From this perspective, human development follows alternative life-history strategies in response to environmental variability, rather than following one species-typical pattern of development.

EDP is closely linked to the theoretical framework of evolutionary psychology (EP), but is also distinct from EP in several domains, including research emphasis (EDP focuses on adaptations of ontogeny, as opposed to adaptations of adulthood) and consideration of proximate ontogenetic and environmental factors (i.e., how development happens) in addition to more ultimate factors (i.e., why development happens), which are the focus of mainstream evolutionary psychology.

Attachment theory

Attachment theory, originally developed by John Bowlby, focuses on the importance of open, intimate, emotionally meaningful relationships. Attachment is described as a biological system or powerful survival impulse that evolved to ensure the survival of the infant. A child who is threatened or stressed will move toward caregivers who create a sense of physical, emotional and psychological safety for the individual. Attachment feeds on body contact and familiarity. Later Mary Ainsworth developed the Strange Situation protocol and the concept of the secure base.

Theorists have proposed four types of attachment styles: secure, anxious-avoidant, anxious-resistant, and disorganized. Secure attachment is a healthy attachment between the infant and the caregiver. It is characterized by trust. Anxious-avoidant is an insecure attachment between an infant and a caregiver. This is characterized by the infant's indifference toward the caregiver. Anxious-resistant is an insecure attachment between the infant and the caregiver characterized by distress from the infant when separated and anger when reunited. Disorganized is an attachment style without a consistent pattern of responses upon return of the parent.

A child can be hindered in its natural tendency to form attachments. Some babies are raised without the stimulation and attention of a regular caregiver or locked away under conditions of abuse or extreme neglect. The possible short-term effects of this deprivation are anger, despair, detachment, and temporary delay in intellectual development. Long-term effects include increased aggression, clinging behavior, detachment, psychosomatic disorders, and an increased risk of depression as an adult.

Attachment style can affect the relationships between people. Attachment is established in early childhood and attachment continues into adulthood. An example of secure attachment continuing in adulthood would be when the person feels confident and is able to meet their own needs. An example of anxious attachment during adulthood is when the adult chooses a partner with anxious-avoidant attachment.

Major debates

Nature vs nurture

A significant issue in developmental psychology is the relationship between innateness and environmental influence in regard to any particular aspect of development. This is often referred to as "nature and nurture" or nativism versus empiricism. A nativist account of development would argue that the processes in question are innate, that is, they are specified by the organism's genes.

An empiricist perspective would argue that those processes are acquired in interaction with the environment. Today developmental psychologists rarely take such polarised positions with regard to most aspects of development; rather they investigate, among many other things, the relationship between innate and environmental influences. One of the ways this relationship has been explored in recent years is through the emerging field of evolutionary developmental psychology.

One area where this innateness debate has been prominently portrayed is in research on language acquisition. A major question in this area is whether or not certain properties of human language are specified genetically or can be acquired through learning. The empiricist position on the issue of language acquisition suggests that the language input provides the necessary information required for learning the structure of language and that infants acquire language through a process of statistical learning. From this perspective, language can be acquired via general learning methods that also apply to other aspects of development, such as perceptual learning.

The nativist position argues that the input from language is too impoverished for infants and children to acquire the structure of language. Linguist Noam Chomsky asserts that, evidenced by the lack of sufficient information in the language input, there is a universal grammar that applies to all human languages and is pre-specified. This has led to the idea that there is a special cognitive module suited for learning language, often called the language acquisition device. Chomsky's critique of the behaviorist model of language acquisition is regarded by many as a key turning point in the decline in the prominence of the theory of behaviorism generally. But Skinner's conception of "Verbal Behavior" has not died, perhaps in part because it has generated successful practical applications.

Continuity vs discontinuity

Since theorists believe that development is a smooth, continuous process, individuals gradually add more of the same types of skills throughout their lives. Other theorists, however, think that development takes place in discontinuous stages. People change rapidly and step up to a new level, and then change very little for a while. With each new step, the person shows interest and responds to the world qualitatively.

Stability vs change

This issue involves the degree to which we become older renditions of our early experience or whether we develop into something different from who we were at an earlier point in development. It considers the extent to which early experiences ( especially infancy) or later experiences are the key determinants of a person's development.

Most lifespan developmentalists, recognise that extreme positions are unwise. Therefore, the key to a comprehensive understanding of development at any stage requires the ·interaction of different factors and not only one.

Mechanisms of Development

Developmental psychology is concerned not only with describing the characteristics of psychological change over time but also seeks to explain the principles and internal workings underlying these changes. Psychologists have attempted to better understand these factors by using models. Developmental models are sometimes computational, but they do not need to be.

A model must simply account for the means by which a process takes place. This is sometimes done in reference to changes in the brain that may correspond to changes in behavior over the course of the development. Computational accounts of development often use either symbolic, connectionist (neural network), or dynamical systems models to explain the Mechanisms of Development.

Research areas

Cognitive development

Cognitive development is primarily concerned with the ways that infants and children acquire, develop, and use internal mental capabilities such as: problem-solving, memory, and language. Major topics in cognitive development are the study of language acquisition and the development of perceptual and motor skills. Piaget was one of the influential early psychologists to study the development of cognitive abilities. His theory suggests that development proceeds through a set of stages from infancy to adulthood and that there is an end point or goal.

Other accounts, such as that of Lev Vygotsky, have suggested that development does not progress through stages, but rather that the developmental process that begins at birth and continues until death is too complex for such structure and finality. Rather, from this viewpoint, developmental processes proceed more continuously. Thus, development should be analyzed, instead of treated as a product to obtain.

K. Warner Schaie has expanded the study of cognitive development into adulthood. Rather than being stable from adolescence, Schaie sees adults as progressing in the application of their cognitive abilities.

Modern cognitive development has integrated the considerations of cognitive psychology and the psychology of individual differences into the interpretation and modeling of development. Specifically, the neo-Piagetian theories of cognitive development showed that the successive levels or stages of cognitive development are associated with increasing processing efficiency and working memory capacity. These increases explain differences between stages, progression to higher stages, and individual differences of children who are the same-age and of the same grade-level. However, other theories have moved away from Piagetian stage theories, and are influenced by accounts of domain-specific information processing, which posit that development is guided by innate evolutionarily-specified and content-specific information processing mechanisms.

Social and emotional development

Developmental psychologists who are interested in social development examine how individuals develop social and emotional competencies. For example, they study how children form friendships, how they understand and deal with emotions, and how identity develops. Research in this area may involve study of the relationship between cognition or cognitive development and social behavior.
Emotional regulation or ER refers to an individual's ability to modulate emotional responses across a variety of contexts. In young children, this modulation is in part controlled externally, by parents and other authority figures. As children develop, they take on more and more responsibility for their internal state. Studies have shown that the development of ER is affected by the emotional regulation children observe in parents and caretakers, the emotional climate in the home, and the reaction of parents and caretakers to the child's emotions.

Music also has an influence on stimulating and enhancing the senses of a child through self-expression.

A child's social and emotional development can be disrupted by motor coordination problems as evidenced by the environmental stress hypothesis. The environmental hypothesis explains how children with coordination problems and developmental coordination disorder are exposed to several psychosocial consequences which act as secondary stressors, leading to an increase in internalizing symptoms such as depression and anxiety. Motor coordination problems affect fine and gross motor movement as well as perceptual-motor skills. Secondary stressors commonly identified include the tendency for children with poor motor skills to be less likely to participate in organized play with other children and more likely to feel socially isolated.

Social and emotional development focuses on 5 keys areas: Self-Awareness, Self Management, Social Awareness, Relationship Skills and Responsible Decision Making.

Physical development

Physical development concerns the physical maturation of an individual's body until it reaches the adult stature. Although physical growth is a highly regular process, all children differ tremendously in the timing of their growth spurts. Studies are being done to analyze how the differences in these timings affect and are related to other variables of developmental psychology such as information processing speed. Traditional measures of physical maturity using x-rays are less in practice nowadays, compared to simple measurements of body parts such as height, weight, head circumference, and arm span.

A few other studies and practices with physical developmental psychology are the phonological abilities of mature 5- to 11-year-olds, and the controversial hypotheses of left-handers being maturationally delayed compared to right-handers. A study by Eaton, Chipperfield, Ritchot, and Kostiuk in 1996 found in three different samples that there was no difference between right- and left-handers.

Memory development

Researchers interested in memory development look at the way our memory develops from childhood and onward. According to Fuzzy-trace theory, we have two separate memory processes: verbatim and gist. These two traces begin to develop at different times as well as at a different pace. Children as young as 4 years-old have verbatim memory, memory for surface information, which increases up to early adulthood, at which point it begins to decline. On the other hand, our capacity for gist memory, memory for semantic information, increases up to early adulthood, at which point it is consistent through old age. Furthermore, our reliance on gist memory traces increases as we age.

Research methods and designs

Main research methods

Developmental psychology employs many of the research methods used in other areas of psychology. However, infants and children cannot be tested in the same ways as adults, so different methods are often used to study their development.

Developmental psychologists have a number of methods to study changes in individuals over time. Common research methods include systematic observation, including naturalistic observation or structured observation; self-reports, which could be clinical interviews or structured interviews; clinical or case study method; and ethnography or participant observation. These methods differ in the extent of control researchers impose on study conditions, and how they construct ideas about which variables to study. Every developmental investigation can be characterized in terms of whether its underlying strategy involves the experimental, correlational, or case study approach. The experimental method involves "actual manipulation of various treatments, circumstances, or events to which the participant or subject is exposed; the experimental design points to cause-and-effect relationships. This method allows for strong inferences to be made of causal relationships between the manipulation of one or more independent variables and subsequent behavior, as measured by the dependent variable. The advantage of using this research method is that it permits determination of cause-and-effect relationships among variables. On the other hand, the limitation is that data obtained in an artificial environment may lack generalizability. The correlational method explores the relationship between two or more events by gathering information about these variables without researcher intervention. The advantage of using a correlational design is that it estimates the strength and direction of relationships among variables in the natural environment; however, the limitation is that it does not permit determination of cause-and-effect relationships among variables. The case study approach allows investigations to obtain an in-depth understanding of an individual participant by collecting data based on interviews, structured questionnaires, observations, and test scores. Each of these methods have its strengths and weaknesses but the experimental method when appropriate is the preferred method of developmental scientists because it provides a controlled situation and conclusions to be drawn about cause-and-effect relationships.

Research designs

Most developmental studies, regardless of whether they employ the experimental, correlational, or case study method, can also be constructed using research designs. Research designs are logical frameworks used to make key comparisons within research studies such as:
In a longitudinal study, a researcher observes many individuals born at or around the same time (a cohort) and carries out new observations as members of the cohort age. This method can be used to draw conclusions about which types of development are universal (or normative) and occur in most members of a cohort. As an example a longitudinal study of early literacy development examined in detail the early literacy experiences of one child in each of 30 families.

Researchers may also observe ways that development varies between individuals, and hypothesize about the causes of variation in their data. Longitudinal studies often require large amounts of time and funding, making them unfeasible in some situations. Also, because members of a cohort all experience historical events unique to their generation, apparently normative developmental trends may, in fact, be universal only to their cohort.

In a cross-sectional study, a researcher observes differences between individuals of different ages at the same time. This generally requires fewer resources than the longitudinal method, and because the individuals come from different cohorts, shared historical events are not so much of a confounding factor. By the same token, however, cross-sectional research may not be the most effective way to study differences between participants, as these differences may result not from their different ages but from their exposure to different historical events.

A third study design, the sequential design, combines both methodologies. Here, a researcher observes members of different birth cohorts at the same time, and then tracks all participants over time, charting changes in the groups. While much more resource-intensive, the format aids in a clearer distinction between what changes can be attributed to an individual or historical environment from those that are truly universal.

Because every method has some weaknesses, developmental psychologists rarely rely on one study or even one method to reach conclusions by finding consistent evidence from as many converging sources as possible.

Life stages of psychological development

Prenatal development

Prenatal development is of interest to psychologists investigating the context of early psychological development. The whole prenatal development involves three main stages: germinal stage, embryonic stage and fetal stage. Germinal stage begins at conception until 2 weeks; embryonic stage means the development from 2 weeks to 8 weeks; fetal stage represents 9 weeks until birth of the baby. The senses develop in the womb itself: a fetus can both see and hear by the second trimester (13 to 24 weeks of age). The sense of touch develops in the embryonic stage (5 to 8 weeks). Most of the brain's billions of neurons also are developed by the second trimester. Babies are hence born with some odor, taste and sound preferences, largely related to the mother's environment.

Some primitive reflexes too arise before birth and are still present in newborns. One hypothesis is that these reflexes are vestigial and have limited use in early human life. Piaget's theory of cognitive development suggested that some early reflexes are building blocks for infant sensorimotor development. For example, the tonic neck reflex may help development by bringing objects into the infant's field of view.

Other reflexes, such as the walking reflex appear to be replaced by more sophisticated voluntary control later in infancy. This may be because the infant gains too much weight after birth to be strong enough to use the reflex, or because the reflex and subsequent development are functionally different. It has also been suggested that some reflexes (for example the moro and walking reflexes) are predominantly adaptations to life in the womb with little connection to early infant development. Primitive reflexes reappear in adults under certain conditions, such as neurological conditions like dementia or traumatic lesions.

Ultrasound has shown that infants are capable of a range of movements in the womb, many of which appear to be more than simple reflexes. By the time they are born, infants can recognize and have a preference for their mother's voice suggesting some prenatal development of auditory perception. Prenatal development and birth complications may also be connected to neurodevelopmental disorders, for example in schizophrenia. With the advent of cognitive neuroscience, embryology and the neuroscience of prenatal development is of increasing interest to developmental psychology research.

Several environmental agents—teratogens—can cause damage during the prenatal period. These include prescription and nonprescription drugs, illegal drugs, tobacco, alcohol, environmental pollutants, infectious disease agents such as the rubella virus and the toxoplasmosis parasite, maternal malnutrition, maternal emotional stress, and Rh factor blood incompatibility between mother and child. There are many statistics which prove the effects of the aforementioned substances. A leading example of this would be that, in America alone, approximately 100,000-375,000 'cocaine babies' are born on an annual basis. This is a result of an expectant mother abusing the drug while pregnant. 'Cocaine babies' are proven to have quite severe and lasting difficulties which persist throughout infancy and right throughout childhood. The drug also encourages behavioural problems in the affected children, as well as defects of various vital organs.

Infancy

From birth until the first year, the child is referred to as an infant. Developmental psychologists vary widely in their assessment of infant psychology, and the influence the outside world has upon it, but certain aspects are relatively clear.

The majority of a newborn infant's time is spent in sleep. At first, this sleep is evenly spread throughout the day and night, but after a couple of months, infants generally become diurnal.

Infants can be seen to have six states, grouped into pairs:
  • quiet sleep and active sleep (dreaming, when REM sleep occurs)
  • quiet waking, and active waking
  • fussing and crying

Infant perception

Infant perception is what a newborn can see, hear, smell, taste, and touch. These five features are better known as one's "five senses". Infants respond to stimuli differently in these different states.
  • Vision is significantly worse in infants than in older children. Infant sight tends to be blurry in early stages but improves over time. Color perception similar to that seen in adults has been demonstrated in infants as young as four months, using habituation methods. Infants get to adult-like vision in about six months.
  • Hearing is well-developed prior to birth, unlike vision. Newborns prefer complex sounds to pure tones, human speech to other sounds, mother's voice to other voices, and the native language to other languages. Scientist believe these features are probably learned in the womb. Infants are fairly good at detecting the direction a sound comes from, and by 18 months their hearing ability is approximately equal to an adult's.
  • Smell and taste are present, with infants showing different expressions of disgust or pleasure when presented with pleasant odors (honey, milk, etc.) or unpleasant odors (rotten egg) and tastes (e.g. sour taste). Newborns are born with odor and taste preferences acquired in the womb from the smell and taste of amniotic fluid, in turn influenced by what the mother eats. Both breast- and bottle-fed babies around 3 days old prefer the smell of human milk to that of formula, indicating an innate preference. There is good evidence for older infants preferring the smell of their mother to that of others.
  • Touch and feel is one of the better-developed senses at birth considering it's one of the first senses to develop inside the womb. This is evidenced by the primitive reflexes described above, and the relatively advanced development of the somatosensory cortex.
  • Pain: Infants feel pain similarly, if not more strongly than older children but pain-relief in infants has not received so much attention as an area of research. Glucose is known to relieve pain in newborns.

Language

Babies are born with the ability to discriminate virtually all sounds of all human languages. Infants of around six months can differentiate between phonemes in their own language, but not between similar phonemes in another language. At this stage infants also start to babble, producing phonemes.

Infant cognition: the Piagetian era

Piaget suggested that an infant's perception and understanding of the world depended on their motor development, which was required for the infant to link visual, tactile and motor representations of objects. According to this view, it is through touching and handling objects that infants develop object permanence, the understanding that objects are solid, permanent, and continue to exist when out of sight.

Special methods are used in the psychological study of infants

Piaget's sensorimotor stage comprised six sub-stages (see sensorimotor stages for more detail). In the early stages, development arises out of movements caused by primitive reflexes. Discovery of new behaviors results from classical and operant conditioning, and the formation of habits. From eight months the infant is able to uncover a hidden object but will persevere when the object is moved.

Piaget came to his conclusion that infants lacked a complete understanding of object permanence before 18 months after observing infants' failure before this age to look for an object where it was last seen. Instead, infants continue to look for an object where it was first seen, committing the "A-not-B error." Some researchers have suggested that before the age of eight to nine months, infants' inability to understand object permanence extends to people, which explains why infants at this age do not cry when their mothers are gone ("Out of sight, out of mind").

Recent findings in infant cognition

In the 1980s and 1990s, researchers have developed many new methods of assessing infants' understanding of the world with far more precision and subtlety than Piaget was able to do in his time. Since then, many studies based on these methods suggest that young infants understand far more about the world than first thought.

Based on recent findings, some researchers (such as Elizabeth Spelke and Renee Baillargeon) have proposed that an understanding of object permanence is not learned at all, but rather comprises part of the innate cognitive capacities of our species.

Other research has suggested that young infants in their first six months of life may possess an understanding of numerous aspects of the world around them, including:
  • an early numerical cognition, that is, an ability to represent number and even compute the outcomes of addition and subtraction operations;
  • an ability to infer the goals of people in their environment;
  • an ability to engage in simple causal reasoning.

Critical periods of development

There are critical periods in infancy and childhood during which development of certain perceptual, sensorimotor, social and language systems depends crucially on environmental stimulation. Feral children such as Genie, deprived of adequate stimulation, fail to acquire important skills and are unable to learn in later childhood. The concept of critical periods is also well-established in neurophysiology, from the work of Hubel and Wiesel among others.

Developmental delays

Children with developmental delays (DD) are at heightened risk for developing clinically significant behavioral and emotional difficulties as compared to children with typical development (TD). However, nearly all studies comparing psychopathology in youth with DD employ TD control groups of the same chronological age (CA).This comorbidity of DD and a mental disorder is often referred to as dual diagnosis. Epidemiological studies indicate that 30–50% of youth with DD meet the clinical cutoff for behavioral and emotional problems and/or diagnosable mental disorder. Studies that include comparison samples of children with typical development (TD) highlight the considerable difference in risk for psychopathology, with the relative risk for youth with DD (to youth with TD) ranging from 2.8–4.1 to 1.

Toddler-hood

Infants shift between ages of one and two to a developmental stage known as toddlerhood. In this stage, an infant's transition into toddlerhood is highlighted through self-awareness, developing maturity in language use, and presence of memory and imagination.

During toddlerhood, babies begin learning how to walk, talk, and make decisions for themselves. An important characteristic of this age period is the development of language, where children are learning how to communicate and express their emotions and desires through the use of vocal sounds, babbling, and eventually words. Self-control also begins to develop. At this age, children take initiative to explore, experiment and learn from making mistakes. Caretakers who encourage toddlers to try new things and test their limits, help the child become autonomous, self-reliant, and confident. If the caretaker is overprotective or disapproving of independent actions, the toddler may begin to doubt their abilities and feel ashamed of the desire for independence. The child's autonomic development is inhibited, leaving them less prepared to deal with the world in the future. Toddlers also begin to identify themselves in gender roles, acting according to their perception of what a man or woman should do.

Socially, the period of toddler-hood is commonly called the "terrible twos". Toddlers often use their new-found language abilities to voice their desires, but are often misunderstood by parents due to their language skills just beginning to develop. A person at this stage testing their independence is another reason behind the stage's infamous label. Tantrums in a fit of frustration are also common.

Childhood

Erik Erikson divides childhood into four stages, each with its distinct social crisis:
  • Stage 1: Infancy (0 to 1½) in which the psychosocial crisis is Trust vs. Mistrust
  • Stage 2: Early childhood (2½ to 3) in which the psychosocial crisis is Autonomy vs. Shame and doubt
  • Stage 3: Play age (3 to 5) in which the psychosocial crisis is Initiative vs. Guilt. (This stage is also called the "pre-school age," "exploratory age" and "toy age.")
  • Stage 4: School age (5 to 12) in which the psychosocial crisis is Industry vs. Inferiority
Play (or preschool) ages 3–5.

In the earliest years, children are "completely dependent on the care of others." Therefore, they develop a "social relationship" with their care givers and, later, with family members. During their preschool years (3-5), they "enlarge their social horizons" to include people outside the family.
Preoperational and then operational thinking develops, which means actions are reversible, and egocentric thought diminishes.

The motor skills of preschoolers increase so they can do more things for themselves. They become more independent. No longer completely dependent on the care of others, the world of this age group expands. More people have a role in shaping their individual personalities. Preschoolers explore and question their world. For Jean Piaget, the child is "a little scientist exploring and reflecting on these explorations to increase competence" and this is done in "a very independent way."

Play is a major activity for ages 3–5. For Piaget, through play "a child reaches higher levels of cognitive development."

In their expanded world, children in the 3-5 age group attempt to find their own way. If this is done in a socially acceptable way, the child develops the initiative. If not, the child develops guilt. Children who develop "guilt" rather than "initiative" have failed Erikson's psychosocial crisis for the 3-5 age group.

Middle childhood ages 6–12.

For Erik Erikson, the psychosocial crisis during middle childhood is Industry vs. Inferiority which, if successfully met, instills a sense of Competency in the child.

In all cultures, middle childhood is a time for developing "skills that will be needed in their society." School offers an arena in which children can gain a view of themselves as "industrious (and worthy)." They are "graded for their school work and often for their industry." They can also develop industry outside of school in sports, games, and doing volunteer work. Children who achieve "success in school or games might develop a feeling of competence."

The "peril during this period is that feelings of inadequacy and inferiority will develop. Parents and teachers can "undermine" a child's development by failing to recognize accomplishments or being overly critical of a child's efforts. Children who are "encouraged and praised" develop a belief in their competence. Lack of encouragement or ability to excel lead to "feelings of inadequacy and inferiority".

The Centers for Disease Control (the CDC) divides Middle Childhood into two stages, 6–8 years and 9–11 years, and gives "developmental milestones for each stage."

Middle Childhood (7-10).

Entering elementary school, children in this age group begin to thinks about the future and their "place in the world." Working with other students and wanting their friendship and acceptance become more important. This leads to "more independence from parents and family." As students, they develop the mental and verbal skills "to describe experiences and talk about thoughts and feelings". They become less self-centered and show "more concern for others".

Middle Childhood (9-11).

For children ages 9–11 "friendships and peer relationships" increase in strength, complexity, and importance. This results in greater "peer pressure." They grow even less dependent on their families and they are challenged academically. To meet this challenge, they increase their attention span and learn to see other points of view.

Adolescence

Adolescence is the period of life between the onset of puberty and the full commitment to an adult social role, such as worker, parent, and/or citizen. It is the period known for the formation of personal and social identity (see Erik Erikson) and the discovery of moral purpose (see William Damon). Intelligence is demonstrated through the logical use of symbols related to abstract concepts and formal reasoning. A return to egocentric thought often occurs early in the period. Only 35% develop the capacity to reason formally during adolescence or adulthood. (Huitt, W. and Hummel, J. January 1998)

It is divided into three parts, namely:
  1. Early Adolescence: 9 to 13 years
  2. Mid Adolescence: 13 to 15 years and
  3. Late Adolescence: 15 to 18 years
The adolescent unconsciously explores questions such as "Who am I? Who do I want to be?" Like toddlers, adolescents must explore, test limits, become autonomous, and commit to an identity, or sense of self. Different roles, behaviors and ideologies must be tried out to select an identity. Role confusion and inability to choose vocation can result from a failure to achieve a sense of identity through, for example, friends.

Early adulthood

Early adulthood generally refers to the period between ages 18 to 25, and according to theorists such as Erik Erikson, is a stage where development is mainly focused on maintaining relationships. Examples include creating bond of intimacy, sustaining friendships, and ultimately making a family. Some theorists state that development of intimacy skills rely on the resolution of previous developmental stages. A sense of identity gained in the previous stages is also necessary for intimacy to develop. If this skill is not learned the alternative is alienation, isolation, a fear of commitment, and the inability to depend on others.

A related framework for studying this part of the lifespan is that of emerging adulthood. Scholars of emerging adulthood, such as Jeffrey Arnett, are not necessarily interested in relationship development. Instead, this concept suggests that people transition after their teenage years into a period not characterized as relationship building and an overall sense of constancy with life, but with years of living with parents, phases of self-discovery, and experimentation.

Middle adulthood

Middle adulthood generally refers to the period between ages 25 to 69. During this period, middle-aged adults experience a conflict between generativity and stagnation. They may either feel a sense of contributing to society, the next generation, or their immediate community; or develop a sense of purposelessness.

Physically, the middle-aged experience a decline in muscular strength, reaction time, sensory keenness, and cardiac output. Also, women experience menopause and a sharp drop in the hormone estrogen. Men experience an equivalent endocrine system event to menopause. Andropause in males is a hormone fluctuation with physical and psychological effects that can be similar to those seen in menopausal females. As men age lowered testosterone levels can contribute to mood swings and a decline in sperm count. Sexual responsiveness can also be affected, including delays in erection and longer periods of penile stimulation required to achieve ejaculation.

Old age

The World Health Organization finds "no general agreement on the age at which a person becomes old." Most "developed countries" set the age as 60 or 65. However, in developing countries inability to make "active contribution" to society, not chronological age, marks the beginning of old age. According to Erikson's stages of psychosocial development, old age is the stage in which individuals assess the quality of their lives. In reflecting on their lives, people in this age group develop a feeling of integrity if deciding that their lives were successful or a feeling of despair if evaluation of one's life indicates a failure to achieve goals.

Physically, older people experience a decline in muscular strength, reaction time, stamina, hearing, distance perception, and the sense of smell. They also are more susceptible to diseases such as cancer and pneumonia due to a weakened immune system. Programs aimed at balance, muscle strength, and mobility have been shown to reduce disability among mildly (but not more severely) disabled elderly.

Sexual expression depends in large part upon the emotional and physical health of the individual. Many older adults continue to be sexually active and satisfied with their sexual activity.

Mental disintegration may also occur, leading to dementia or ailments such as Alzheimer's disease. It is generally believed that crystallized intelligence increases up to old age, while fluid intelligence decreases with age. Whether or not normal intelligence increases or decreases with age depends on the measure and study. Longitudinal studies show that perceptual speed, inductive reasoning, and spatial orientation decline. An article on adult cognitive development reports that cross-sectional studies show that "some abilities remained stable into early old age."

Parenting

Parenting variables alone have typically accounted for 20 to 50 percent of the variance in child outcomes.

All parents have their own parenting styles. Parenting styles, according to Kimberly Kopoko, are "based upon two aspects of parenting behavior; control and warmth. Parental control refers to the degree to which parents manage their children's behavior. Parental warmth refers to the degree to which parents are accepting and responsive to their children's behavior."

Parenting styles

The following parenting styles have been described in the child development literature:
  • Authoritative parenting is characterized as parents who have high parental warmth, responsiveness, and demandingness, but rate low in negativity and conflict. These parents are assertive but not intrusive or overly restrictive. This method of parenting is associated with more positive social and academic outcomes. The beneficial outcomes of authoritative parenting are not necessarily universal. Among African American adolescents, authoritative parenting is not associated with academic achievement without peer support for achievement. Children who are raised by authoritative parents are "more likely to become independent, self-reliant, socially accepted, academically successful, and well-behaved. They are less likely to report depression and anxiety, and less likely to engage in antisocial behavior like delinquency and drug use."
  • Authoritarian parenting is characterized by low levels of warmth and responsiveness with high levels of demandingness and firm control. These parents focus on obedience and they monitor their children regularly. In general, this style of parenting is associated with maladaptive outcomes. The outcomes are more harmful for middle class boys than girls, preschool white girls than preschool black girls, and for white boys than Hispanic boys. Furthermore, the negative effects of authoritarian parenting among Asian Americans can be offset by positive peer support. Finally, among African Americans, some elements of authoritarian parenting such as firm control and physical discipline do not serve as predictive factors for negative outcomes.
  • Permissive parenting is characterized by high levels of responsiveness combined with low levels of demandingness. These parents are lenient and do not necessarily require mature behavior. They allow for a high degree of self-regulation and typically avoid confrontation. Compared to children raised using the authoritative style, preschool girls raised in permissive families are less assertive. Additionally, preschool children of both sexes are less cognitively competent than those children raised under authoritative parenting styles.
  • Rejecting or neglectful parenting is the final category. This is characterized by low levels of demandingness and responsiveness. These parents are typically disengaged in their child's lives, lacking structure in their parenting styles and are unsupportive. Children in this category are typically the least competent of all the categories.

Mother and father factors

Parenting roles in child development have typically focused on the role of the mother. Recent literature, however, has looked toward the father as having an important role in child development. Affirming a role for fathers, studies have shown that children as young as 15 months benefit significantly from substantial engagement with their father. In particular, a study in the U.S. and New Zealand found the presence of the natural father was the most significant factor in reducing rates of early sexual activity and rates of teenage pregnancy in girls. Furthermore, another argument is that neither a mother nor a father is actually essential in successful parenting, and that single parents as well as homosexual couples can support positive child outcomes. According to this set of research, children need at least one consistently responsible adult with whom the child can have a positive emotional connection. Having more than one of these figures contributes to a higher likelihood of positive child outcomes.

Divorce

Another parental factor often debated in terms of its effects on child development is divorce. Divorce in itself is not a determining factor of negative child outcomes. In fact, the majority of children from divorcing families fall into the normal range on measures of psychological and cognitive functioning. A number of mediating factors play a role in determining the effects divorce has on a child, for example, divorcing families with young children often face harsher consequences in terms of demographic, social, and economic changes than do families with older children. Positive coparenting after divorce is part of a pattern associated with positive child coping, while hostile parenting behaviors lead to a destructive pattern leaving children at risk. Additionally, direct parental relationship with the child also affects the development of a child after a divorce. Overall, protective factors facilitating positive child development after a divorce are maternal warmth, positive father-child relationship, and cooperation between parents.

Development of the nervous system


From Wikipedia, the free encyclopedia


Development of the nervous system refers to the processes that generate, shape, and reshape the nervous system of animals, from the earliest stages of embryogenesis to adulthood. The field of neural development draws on both neuroscience and developmental biology to describe and provide insight into the cellular and molecular mechanisms by which complex nervous systems develop, from the nematode and fruit fly to mammals. Defects in neural development can lead to malformations and a wide variety of sensory, motor, and cognitive impairments, including holoprosencephaly and other neurological disorders such as Rett syndrome, Down syndrome and intellectual disability.

Overview of brain development

The mammalian central nervous system (CNS) is derived from the ectoderm—the outermost tissue layer—of the embryo. In the third week of human development the neuroectoderm appears and forms the neural plate along the dorsal side of the embryo. The neural plate is the source of the majority of neurons and glial cells of the CNS. A groove forms along the long axis of the neural plate and, by week four of development, the neural plate wraps in on itself to give rise to the neural tube, which is filled with cerebrospinal fluid (CSF). As the embryo develops, the anterior part of the neural tube forms a series of bulges called vesicles, which become the primary anatomical regions of the brain: the forebrain (prosencephalon), midbrain (mesencephalon), and hindbrain (rhombencephalon). These simple, early vesicles enlarge and further divide into the telencephalon (future cerebral cortex and basal ganglia), diencephalon (future thalamus and hypothalamus), mesencephalon (future colliculi), metencephalon (future pons and cerebellum), and myelencephalon (future medulla). The CSF-filled central chamber is continuous from the telencephalon to the spinal cord, and constitutes the developing ventricular system of the CNS. Because the neural tube gives rise to the brain and spinal cord any mutations at this stage in development can lead to lethal deformities like anencephaly or lifelong disabilities like spina bifida. During this time, the walls of the neural tube contain neural stem cells, which drive brain growth as they divide many times. Gradually some of the cells stop dividing and differentiate into neurons and glial cells, which are the main cellular components of the CNS. The newly generated neurons migrate to different parts of the developing brain to self-organize into different brain structures. Once the neurons have reached their regional positions, they extend axons and dendrites, which allow them to communicate with other neurons via synapses. Synaptic communication between neurons leads to the establishment of functional neural circuits that mediate sensory and motor processing, and underlie behavior.
Flowchart of human brain development

Aspects

Some landmarks of neural development include the birth and differentiation of neurons from stem cell precursors, the migration of immature neurons from their birthplaces in the embryo to their final positions, outgrowth of axons and dendrites from neurons, guidance of the motile growth cone through the embryo towards postsynaptic partners, the generation of synapses between these axons and their postsynaptic partners, and finally the lifelong changes in synapses, which are thought to underlie learning and memory.

Typically, these neurodevelopmental processes can be broadly divided into two classes: activity-independent mechanisms and activity-dependent mechanisms. Activity-independent mechanisms are generally believed to occur as hardwired processes determined by genetic programs played out within individual neurons. These include differentiation, migration and axon guidance to their initial target areas. These processes are thought of as being independent of neural activity and sensory experience. Once axons reach their target areas, activity-dependent mechanisms come into play. Although synapse formation is an activity-independent event, modification of synapses and synapse elimination requires neural activity.

Developmental neuroscience uses a variety of animal models including mice Mus musculus, the fruit fly Drosophila melanogaster, the zebrafish Danio rerio, Xenopus laevis tadpoles and the worm Caenorhabditis elegans, among others.

Myelination, formation of the lipid myelin bilayer around neuronal axons, is a process that is essential for normal brain function. The myelin sheath provides insulation for the nerve impulse when communicating between neural systems. Without it, the impulse would be disrupted and the signal would not reach its target, thus impairing normal functioning. Because so much of brain development occurs in the prenatal stage and infancy, it is crucial that myelination, along with cortical development occur properly. Magnetic resonance imaging (MRI) is a non-invasive technique used to investigate myelination and cortical maturation (the cortex is the outer layer of the brain composed of gray matter). Rather than showing the actual myelin, the MRI picks up on the myelin water fraction (MWF), a measure of myelin content. Multicomponent relaxometry (MCR) allow visualization and quantification of myelin content. MCR is also useful for tracking white matter maturation, which plays an important role in cognitive development. It has been discovered that in infancy, myelination occurs in a posterior-to-anterior pattern. Because there is little evidence of a relationship between myelination and cortical thickness, it was revealed that cortical thickness is independent of white matter MWF. This allows various aspects of the brain to grow simultaneously, leading to a more fully developed brain.

Neural induction

During early embryonic development the ectoderm becomes specified to give rise to the epidermis (skin) and the neural plate. The conversion of undifferentiated ectoderm to neuro-ectoderm requires signals from the mesoderm. At the onset of gastrulation presumptive mesodermal cells move through the dorsal blastopore lip and form a layer in between the endoderm and the ectoderm. These mesodermal cells that migrate along the dorsal midline give rise to a structure called the notochord. Ectodermal cells overlying the notochord develop into the neural plate in response to a diffusible signal produced by the notochord. The remainder of the ectoderm gives rise to the epidermis (skin). The ability of the mesoderm to convert the overlying ectoderm into neural tissue is called neural induction.

The neural plate folds outwards during the third week of gestation to form the neural groove. Beginning in the future neck region, the neural folds of this groove close to create the neural tube. The formation of the neural tube from the ectoderm is called neurulation. The ventral part of the neural tube is called the basal plate; the dorsal part is called the alar plate. The hollow interior is called the neural canal. By the end of the fourth week of gestation, the open ends of the neural tube, called the neuropores, close off.

A transplanted blastopore lip can convert ectoderm into neural tissue and is said to have an inductive effect. Neural inducers are molecules that can induce the expression of neural genes in ectoderm explants without inducing mesodermal genes as well. Neural induction is often studied in xenopus embryos since they have a simple body pattern and there are good markers to distinguish between neural and non-neural tissue. Examples of neural inducers are the molecules noggin and chordin.

When embryonic ectodermal cells are cultured at low density in the absence of mesodermal cells they undergo neural differentiation (express neural genes), suggesting that neural differentiation is the default fate of ectodermal cells. In explant cultures (which allow direct cell-cell interactions) the same cells differentiate into epidermis. This is due to the action of BMP4 (a TGF-β family protein) that induces ectodermal cultures to differentiate into epidermis. During neural induction, noggin and chordin are produced by the dorsal mesoderm (notochord) and diffuse into the overlying ectoderm to inhibit the activity of BMP4. This inhibition of BMP4 causes the cells to differentiate into neural cells. Inhibition of TGF-β and BMP (bone morphogenetic protein) signaling can efficiently induce neural tissue from human pluripotent stem cells, a model of early human development.

Regionalization

Late in the fourth week, the superior part of the neural tube flexes at the level of the future midbrain—the mesencephalon. Above the mesencephalon is the prosencephalon (future forebrain) and beneath it is the rhombencephalon (future hindbrain).

The optical vesicle (which eventually become the optic nerve, retina and iris) forms at the basal plate of the prosencephalon. The alar plate of the prosencephalon expands to form the cerebral hemispheres (the telencephalon) whilst its basal plate becomes the diencephalon. Finally, the optic vesicle grows to form an optic outgrowth.

Patterning of the nervous system

In chordates, dorsal ectoderm forms all neural tissue and the nervous system. Patterning occurs due to specific environmental conditions - different concentrations of signaling molecules

Dorsoventral axis

The ventral half of the neural plate is controlled by the notochord, which acts as the 'organiser'. The dorsal half is controlled by the ectoderm plate, which flanks either side of the neural plate.

Ectoderm follows a default pathway to become neural tissue. Evidence for this comes from single, cultured cells of ectoderm, which go on to form neural tissue. This is postulated to be because of a lack of BMPs, which are blocked by the organiser. The organiser may produce molecules such as follistatin, noggin and chordin that inhibit BMPs.

The ventral neural tube is patterned by sonic hedgehog (Shh) from the notochord, which acts as the inducing tissue. Notochord-derived Shh signals to the floor plate, and induces Shh expression in the floor plate. Floor plate-derived Shh subsequently signals to other cells in the neural tube, and is essential for proper specification of ventral neuron progenitor domains. Loss of Shh from the notochord and/or floor plate prevents proper specification of these progenitor domains. Shh binds Patched1, relieving Patched-mediated inhibition of Smoothened, leading to activation of Gli family of transcription factors (Gli1, Gli2, and Gli3) transcription factors.

In this context Shh acts as a morphogen - it induces cell differentiation dependent on its concentration. At low concentrations it forms ventral interneurones, at higher concentrations it induces motor neuron development, and at highest concentrations it induces floor plate differentiation. Failure of Shh-modulated differentiation causes holoprosencephaly.

The dorsal neural tube is patterned by BMPs from the epidermal ectoderm flanking the neural plate. These induce sensory interneurones by activating Sr/Thr kinases and altering SMAD transcription factor levels.

Rostrocaudal (Anteroposterior) axis

Signals that control anteroposterior neural development include FGF and retinoic acid, which act in the hindbrain and spinal cord. The hindbrain, for example, is patterned by Hox genes, which are expressed in overlapping domains along the anteroposterior axis under the control of retinoic acid. The 3' genes in the Hox cluster are induced by retinoic acid in the hindbrain, whereas the 5' Hox genes are not induced by retinoic acid and are expressed more posteriorly in the spinal cord. Hoxb-1 is expressed in rhombomere 4 and gives rise to the facial nerve. Without this Hoxb-1 expression, a nerve similar to the trigeminal nerve arises.

Neurogenesis

Neurogenesis is the process by which neurons are generated from neural stem cells and progenitor cells. Neurons are 'post-mitotic', meaning that they will never divide again for the lifetime of the organism.

Neuronal migration

Corticogenesis: younger neurons migrate past older ones using radial glia as a scaffolding. Cajal-Retzius cells (red) release reelin (orange)

Neuronal migration is the method by which neurons travel from their origin or birthplace to their final position in the brain. There are several ways they can do this, e.g. by radial migration or tangential migration. This time lapse displays sequences of radial migration (also known as glial guidance) and somal translocation.

Tangential migration of interneurons from ganglionic eminence

Radial migration

Neuronal precursor cells proliferate in the ventricular zone of the developing neocortex, where the principal neural stem cell is the radial glial cell. The first postmitotic cells must leave the stem cell niche and migrate outward to form the preplate, which is destined to become Cajal-Retzius cells and subplate neurons. These cells do so by somal translocation. Neurons migrating with this mode of locomotion are bipolar and attach the leading edge of the process to the pia. The soma is then transported to the pial surface by nucleokinesis, a process by which a microtubule "cage" around the nucleus elongates and contracts in association with the centrosome to guide the nucleus to its final destination. Radial glial cells, whose fibers serve as a scaffolding for migrating cells and a means of radial communication mediated by calcium dynamic activity, act as the main excitatory neuronal stem cell of the cerebral cortex or translocate to the cortical plate and differentiate either into astrocytes or neurons. Somal translocation can occur at any time during development.

Subsequent waves of neurons split the preplate by migrating along radial glial fibres to form the cortical plate. Each wave of migrating cells travel past their predecessors forming layers in an inside-out manner, meaning that the youngest neurons are the closest to the surface. It is estimated that glial guided migration represents 90% of migrating neurons in human and about 75% in rodents.

Tangential migration

Most interneurons migrate tangentially through multiple modes of migration to reach their appropriate location in the cortex. An example of tangential migration is the movement of interneurons from the ganglionic eminence to the cerebral cortex. One example of ongoing tangential migration in a mature organism, observed in some animals, is the rostral migratory stream connecting subventricular zone and olfactory bulb.

Axophilic migration

Many neurons migrating along the anterior-posterior axis of the body use existing axon tracts to migrate along; this is called axophilic migration. An example of this mode of migration is in GnRH-expressing neurons, which make a long journey from their birthplace in the nose, through the forebrain, and into the hypothalamus. Many of the mechanisms of this migration have been worked out, starting with the extracellular guidance cues that trigger intracellular signaling. These intracellular signals, such as calcium signaling, lead to actin  and microtubule cytoskeletal dynamics, which produce cellular forces that interact with the extracellular environment through cell adhesion proteins to cause the movement of these cells.

Other modes of migration

There is also a method of neuronal migration called multipolar migration. This is seen in multipolar cells, which are abundantly present in the cortical intermediate zone. They do not resemble the cells migrating by locomotion or somal translocation. Instead these multipolar cells express neuronal markers and extend multiple thin processes in various directions independently of the radial glial fibers.

Neurotrophic factors

The survival of neurons is regulated by survival factors, called trophic factors. The neurotrophic hypothesis was formulated by Victor Hamburger and Rita Levi Montalcini based on studies of the developing nervous system. Victor Hamburger discovered that implanting an extra limb in the developing chick led to an increase in the number of spinal motor neurons. Initially he thought that the extra limb was inducing proliferation of motor neurons, but he and his colleagues later showed that there was a great deal of motor neuron death during normal development, and the extra limb prevented this cell death. According to the neurotrophic hypothesis, growing axons compete for limiting amounts of target-derived trophic factors and axons that fail to receive sufficient trophic support die by apoptosis. It is now clear that factors produced by a number of sources contribute to neuronal survival.
  • Nerve Growth Factor (NGF): Rita Levi Montalcini and Stanley Cohen purified the first trophic factor, Nerve Growth Factor (NGF), for which they received the Nobel Prize. There are three NGF-related trophic factors: BDNF, NT3, and NT4, which regulate survival of various neuronal populations. The Trk proteins act as receptors for NGF and related factors. Trk is a receptor tyrosine kinase. Trk dimerization and phosphorylation leads to activation of various intracellular signaling pathways including the MAP kinase, Akt, and PKC pathways.
  • CNTF: Ciliary neurotrophic factor is another protein that acts as a survival factor for motor neurons. CNTF acts via a receptor complex that includes CNTFRα, GP130, and LIFRβ. Activation of the receptor leads to phosphorylation and recruitment of the JAK kinase, which in turn phosphorylates LIFRβ. LIFRβ acts as a docking site for the STAT transcription factors. JAK kinase phosphorylates STAT proteins, which dissociate from the receptor and translocate to the nucleus to regulate gene expression.
  • GDNF: Glial derived neurotrophic factor is a member of the TGFb family of proteins, and is a potent trophic factor for striatal neurons. The functional receptor is a heterodimer, composed of type 1 and type 2 receptors. Activation of the type 1 receptor leads to phosphorylation of Smad proteins, which translocate to the nucleus to activate gene expression.

Synapse formation

Neuromuscular junction

Much of our understanding of synapse formation comes from studies at the neuromuscular junction. The transmitter at this synapse is acetylcholine. The acetylcholine receptor (AchR) is present at the surface of muscle cells before synapse formation. The arrival of the nerve induces clustering of the receptors at the synapse. McMahan and Sanes showed that the synaptogenic signal is concentrated at the basal lamina. They also showed that the synaptogenic signal is produced by the nerve, and they identified the factor as Agrin. Agrin induces clustering of AchRs on the muscle surface and synapse formation is disrupted in agrin knockout mice. Agrin transduces the signal via MuSK receptor to rapsyn. Fischbach and colleagues showed that receptor subunits are selectively transcribed from nuclei next to the synaptic site. This is mediated by neuregulins.

In the mature synapse each muscle fiber is innervated by one motor neuron. However, during development many of the fibers are innervated by multiple axons. Lichtman and colleagues have studied the process of synapses elimination. This is an activity-dependent event. Partial blockage of the receptor leads to retraction of corresponding presynaptic terminals.

CNS synapses

Agrin appears not to be a central mediator of CNS synapse formation and there is active interest in identifying signals that mediate CNS synaptogenesis. Neurons in culture develop synapses that are similar to those that form in vivo, suggesting that synaptogenic signals can function properly in vitro. CNS synaptogenesis studies have focused mainly on glutamatergic synapses. Imaging experiments show that dendrites are highly dynamic during development and often initiate contact with axons. This is followed by recruitment of postsynaptic proteins to the site of contact. Stephen Smith and colleagues have shown that contact initiated by dendritic filopodia can develop into synapses.
Induction of synapse formation by glial factors: Barres and colleagues made the observation that factors in glial conditioned media induce synapse formation in retinal ganglion cell cultures. Synapse formation in the CNS is correlated with astrocyte differentiation suggesting that astrocytes might provide a synaptogenic factor. The identity of the astrocytic factors is not yet known.

Neuroligins and SynCAM as synaptogenic signals: Sudhof, Serafini, Scheiffele and colleagues have shown that neuroligins and SynCAM can act as factors that induce presynaptic differentiation. Neuroligins are concentrated at the postsynaptic site and act via neurexins concentrated in the presynaptic axons. SynCAM is a cell adhesion molecule that is present in both pre- and post-synaptic membranes.

Activity dependent mechanisms in the assembly of neural circuits

The processes of neuronal migration, differentiation and axon guidance are generally believed to be activity-independent mechanisms and rely on hard-wired genetic programs in the neurons themselves. New research findings however have implicated a role for activity-dependent mechanisms in mediating some aspects of the aforementioned processes such as the rate of neuronal migration, aspects of neuronal differentiation and axon pathfinding. Activity-dependent mechanisms influence neural circuit development and are crucial for laying out early connectivity maps and the continued refinement of synapses which occurs during development. There are two distinct types of neural activity we observe in developing circuits -early spontaneous activity and sensory-evoked activity. Spontaneous activity occurs early during neural circuit development even when sensory input is absent and is observed in many systems such as the developing visual system, auditory system, motor system, hippocampus, cerebellum, and neocortex.

Experimental techniques such as direct electrophysiological recording, fluorescence imaging using calcium indicators and optogenetic techniques have shed light on the nature and function of these early bursts of activity. They have distinct spatial and temporal patterns during development and their ablation during development has been known to result in deficits in network refinement in the visual system. In the immature retina, waves of spontaneous action potentials arise from the retinal ganglion cells and sweep across the retinal surface in the first few postnatal weeks. These waves are mediated by neurotransmitter acetylcholine in the initial phase and later on by glutamate. They are thought to instruct the formation of two sensory maps- the retinotopic map and eye-specific segregation. Retinotopic map refinement occurs in downstream visual targets in the brain-the superior colliculus (SC) and dorsal lateral geniculate nucleus (LGN). Pharmacological disruption and mouse models lacking the β2 subunit of the nicotinic acetylcholine receptor has shown that the lack of spontaneous activity leads to marked defects in retinotopy and eye-specific segregation.

In the developing auditory system, developing cochlea generate bursts of activity which spreads across the inner hair cells and spiral ganglion neurons which relay auditory information to the brain. ATP release from supporting cells triggers action potentials in inner hair cells. In the auditory system, spontaneous activity is thought to be involved in tonotopic map formation by segregating cochlear neuron axons tuned to high and low frequencies. In the motor system, periodic bursts of spontaneous activity are driven by excitatory GABA and glutamate during the early stages and by acetylcholine and glutamate at later stages. In the developing zebrafish spinal cord, early spontaneous activity is required for the formation of increasingly synchronous alternating bursts between ipsilateral and contralateral regions of the spinal cord and for the integration of new cells into the circuit. In the cortex, early waves of activity have been observed in the cerebellum and cortical slices. Once sensory stimulus becomes available, final fine-tuning of sensory-coding maps and circuit refinement begins to rely more and more on sensory-evoked activity as demonstrated by classic experiments about the effects of sensory deprivation during critical periods.

Contemporary diffusion-weigthted MRI techniques may also uncover the macroscopic process of axonal development. The connectome can be constructed from diffusion MRI data: the vertices of the graph correspond to anatomically labelled gray matter areas, and two such vertices, say u and v, are connected by an edge if the tractography phase of the data processing finds an axonal fiber that connects the two areas, corresponding to u and v.
Consensus Connectome Dynamics

Numerous braingraphs, computed from the Human Connectome Project can be downloaded from the http://braingraph.org site. The Consensus Connectome Dynamics (CCD) is a remarkable phenomenon that was discovered by continuously decreasing the minimum confidence-parameter at the graphical interface of the Budapest Reference Connectome Server. The Budapest Reference Connectome Server (http://connectome.pitgroup.org) depicts the cerebral connections of n=418 subjects with a frequency-parameter k: For any k=1,2,...,n one can view the graph of the edges that are present in at least k connectomes. If parameter k is decreased one-by-one from k=n through k=1 then more and more edges appear in the graph, since the inclusion condition is relaxed. The surprising observation is that the appearance of the edges is far from random: it resembles a growing, complex structure, like a tree or a shrub (visualized on the animation on the left).

It is hypothesized in  that the growing structure copies the axonal development of the human brain: the earliest developing connections (axonal fibers) are common at most of the subjects, and the subsequently developing connections have larger and larger variance, because their variances are accumulated in the process of axonal development.

Synapse elimination

Several motorneurons compete for each neuromuscular junction, but only one survives until adulthood. Competition in vitro has been shown to involve a limited neurotrophic substance that is released, or that neural activity infers advantage to strong post-synaptic connections by giving resistance to a toxin also released upon nerve stimulation. In vivo, it is suggested that muscle fibres select the strongest neuron through a retrograde signal.

Adult neurogenesis

Contrary to popular belief, neurogenesis also occurs in specific parts of the adult brain.

Computer-aided software engineering

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