Brain Development and The Role of Experience in TH
Brain Development and The Role of Experience in TH
Brain Development and The Role of Experience in TH
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T
he past 30 years of research have provided a new and deeper ment can play a role; for example, it
understanding of the brain and its role in psychological is well known that the lack of nutrition
functions. In particular, researchers now have a better sense of (e.g., folic acid) and the presence of toxins
how brain development affects the development of behavior. (e.g., alcohol) can both deleteriously influ-
Measurement techniques such as electroencephalogram (EEG) ence the developing brain. In contrast, much
of brain development that occurs postna-
and event related potentials (ERP) can be used to study infants,
tally is experience-dependent and defined by
children, and adults, and this flexibility has allowed researchers gene–environment interactions. Below we
to investigate a variety of developmental processes. provide brief descriptions of the anatomical
changes that characterize the early stages of
Research using these measures on the on experience has helped refine researchers’ brain development.
developing brain has clarified several argu- understanding of how developmental pro-
ments about the nature of child development cesses are fueled. Neurulation
and informed debates such as those sur- In the following paragraphs, we will exam- About 2 weeks after conception, the devel-
rounding the state of the infant’s brain at ine some of the essential ideas that have oping embryo has organized itself into a
birth (whether it is a “blank slate” or not), the helped researchers understand the develop- three-layered, spherical structure. In one area
identification of critical periods of develop- ment of the human brain in the early years of
ment, and the relative importance of genes life. We begin with an overview of the stages
versus environment. involved in the anatomical development of
It is important to note that, although the brain. Subsequently, we examine three
much of the research has been conducted topics that research in brain development Abstract
on infants, it is a collaborative effort has uncovered, clarified, and elaborated: how Research over the past several
between infant and animal research that development is hierarchically structured, decades has provided insight into the
has uncovered the neurobiological princi- such that later development depends on early processes that govern early brain
ples that govern development in humans. development; how experience in the first year development and how those processes
Researchers have made use of the homology of life modulates the plasticity of the brain; contribute to behavior. In the following
that exists among developing nervous sys- and how early deprivation has strong and last- article, we provide an overview of early
tems of different species, and many of the ing effects on the brain. brain development beginning with a
cutting-edge ideas discussed in the develop- summary of the prenatal period. We
mental literature have their origins in animal Early Stages of Brain Development then turn to postnatal development
research—but they have been tested and
clarified in neurobehavioral experiments
with infants and young children. In humans,
researchers can investigate the neural corre-
lates of behavior whereas in animals they can
A n account of brain development in
the early years of childhood is only
complete if we first examine the ori-
gins of this process during the prenatal
months. Brain development is a protracted
and examine how brain functions are
built and how experience mediates
this process. Specifically, we discuss
findings from research on speech and
on face processing. The results of
dig deeper into the mechanisms that drive the process that begins about 2 weeks after con- this research highlight how the first
processes that these neural correlates reflect. ception and continues into young adulthood few years of life are a particularly
To this end, much of human brain research 20 years later. Brain development that occurs important period of development of the
in the past three decades has focused on the during the prenatal months is largely under brain.
brain basis of behavior. A more recent a focus genetic control, although clearly the environ-
After the cells are born, they travel to their for brain function.
final destinations. The cerebral cortex is
composed of multilayered tissue several mil- Synaptogenesis
limeters thick. It is formed by the movement A synapse is a point of contact between
of cells in an inside-out direction, begin- two brain cells, often two neurons and fre-
ning in the ventricular zone and migrating quently a dendrite and an axon. The first
through the intermediate zone, with the cells synapses are generally observed by about the
eventually reaching their final destination 23rd week of gestation (Molliver, Kostovic,
on the outside of the developing brain. The & Van der Loos, 1973), although the peak of
earliest migrating cells occupy the deepest production does not occur until some time
cortical layer, whereas the subsequent migra- in the first year of life. As is the case with
Synapse reduction, or pruning, is highly tions pass through previously formed layers neurons, massive overproduction of syn-
dependent on experience and serves as to form the outer layers. About 25 weeks after apses is followed by a gradual reduction.
the basis of much of the learning that conception, all six layers of the cortex will This process of synapse reduction, or prun-
occurs during the early years of life. have formed. ing, is highly dependent on experience and
The inside-out pattern of migration serves as the basis of much of the learning
described here is that of radial migration, that occurs during the early years of life. It
of this sphere, the cells thicken to form what which applies to about 70%–80% of migrating is important to note that the various struc-
is called the neural plate. This plate then folds neurons, most of which are pyramidal tures of the brain reach their peak of synapse
over onto itself, forming a tube that gradu- neurons and glia. Pyramidal neurons are production at different points. In the visual
ally closes first at the bottom and then at the the large neurons in the cortex that are cortex, for example, the peak is reached
top, much like a zipper. This creates the neu- responsible for sending signals to different somewhere between the 4th and 8th postna-
ral tube, the inner cells of which will lead to layers of the cortex and other parts of the tal month, but areas of the prefrontal cortex
the formation of the central nervous sys- brain. Glia are nonneuronal brain cells that do not reach their peak until the 15th postna-
tem (brain and spinal cord) while the outer are involved in the support of neuronal tal month. The difference in timing in peak
cells will give rise to the autonomic nervous processes (such as producing myelin or synapse production is important because it
system (nerves outside the brain and spinal removing debris, including dead brain cells). affects the timing of the plasticity of these
cord). In contrast, interneurons—relatively smaller regions; the later the peak synapse produc-
Once the neural tube is closed, it neurons that are involved in communication tion, the longer the region remains plastic.
becomes a three-vesicle structure and between pyramidal cells within a particular
shortly thereafter a five-vesicle structure. layer of the cortex—follow a pattern of Synapse Pruning
The different regions of tissue around tangential migration. The overproduction of synapses is fol-
the ventricles will become distinct brain lowed by a pruning back of the unused and
structures. The anterior portion of the tube Differentiation overabundance of synapses. Until the stage
will become the forebrain, which includes Once a neuron has migrated to its target of synaptogenesis, the stages of brain devel-
the cerebral hemispheres; the diencephalon destination, it generally proceeds along one opment are largely gene driven. However,
(the thalamus and the hypothalamus); of two roads: It can differentiate into a mature once the brain reaches the point where syn-
and the basal ganglia. The cells around the neuron, complete with an axon and dendrites, apses are eliminated, the balance shifts;
middle vesicle will become the midbrain, a or it can be retracted through apoptosis. the process of pruning is largely experience
structure that connects the diencephalon to Current estimates suggest that the number driven. As with synapse production, the tim-
the hindbrain. The rear-most portion of the of neurons that are retracted is between 40% ing of synapse pruning is dependent on the
tube will give rise to the hindbrain, which will and 60% (Oppenheim & Johnson, 2003). The area of the brain in which it occurs. In the
consist of the medulla oblongata, the pons, development of axons is facilitated by growth parts of the cortex involved in visual and
and the cerebellum. Finally, the cells that cones, small structures that form at the edge auditory perception, for example, pruning
remain will give rise to the spinal cord. of an axon. The cellular processes that occur is complete between the 4th and 6th year of
at the growth cone promote growth toward life. In contrast, pruning in areas involved in
Proliferation certain targets and away from others. Such higher cognitive functions (such as inhibitory
Once the general structure of the neural processes are driven by molecular guidance control and emotion regulation) contin-
tube has been laid out, the cells that line the cues as well as by anatomic structures at the ues through adolescence (Huttenlocher &
innermost part of the tube, called the ventric- tip of the growth cone. Dabholkar, 1997). The processes of overpro-
ular zone, proliferate at a logarithmic rate. As Dendrite formation occurs by a slightly duction of synapses and subsequent synaptic
these cells multiply, they form a second zone, different process, one that is thought to be reduction are essential for the flexibility
T
respond to the unique environment in which a period particularly important to develop- he brain is much more sensitive to
he or she is born. Those pathways that are ment of a healthy brain. The foundations of experience in the first few years of
activated by the environment are strength- sensory and perceptual systems that are crit- life than in later years. The plasticity
ened while the ones that go unused are ical to language, social behavior, and emotion of the brain underlies much of the learning
eliminated. In this way, the networks of neu- are formed in the early years and are strongly that occurs during this period. In the lan-
rons involved in the development of behavior influenced by experiences during this time. guage example in the previous section, we
are fine-tuned and modified as needed. This is not to say that later development can- noted that infants are sensitive to most lan-
not affect these behaviors—on the contrary, guage sounds in the first half-year of life but
Myelination experiences later in life are also very impor- during the second half they begin to special-
The final process involved in the develop- tant to the function of the brain. However, ize in their native tongue at the expense of
ment of the brain is called myelination. In this experiences in the early years of childhood the broad sensitivity to nonnative language
process the axons of neurons are wrapped in affect the development of brain architec- sounds. The period of heightened sensitivity
fatty cells, which ultimately facilitates neu- ture in a way that later experiences do not. In to language exposure is not, however, a crit-
ronal activity and communication because the following pages we will elaborate on how ical period in the sense that infants can no
this insulation allows myelinated axons to experience affects development between longer learn the sounds of another language
transmit electrical signals faster than unmy- birth and 3 years of age. once it is over. In fact, 12-month-old infants
elinated axons. The timing of myelination given additional experience with speech
is dependent on the region of the brain in Brain Beginnings: Constructing a sounds from a nonnative language continue
which it occurs. Regions of the brain in cer- Foundation for the Future to be able to discriminate among sounds
tain sensory and motor areas are myelinated
earlier in a process that is complete around
the preschool period. In contrast, in regions
involved in higher cognitive abilities, such as
the prefrontal cortex, the process is not com-
T he development of the brain is a life-
long process. Indeed, recent research
suggests that the brain is capable of
changing throughout the lifespan (Crawford,
Pesch, & von Noorden, 1996; Jones, 2000;
(Kuhl et al., 2003).
Similarly, in the domain of face pro-
cessing, an index of development of visual
perception important to social behavior,
6-month-olds, 9-month-olds, and adults are
plete until adolescence or early adulthood Keuroghlian & Knudsen, 2007), although all equally capable of discriminating between
(for recent reviews see Nelson, de Haan, & perhaps not in all ways (e.g., humans do not two human faces, whereas 6-month-olds
Thomas, 2006; Nelson & Jeste, 2008). “learn” to see or hear better as they age). alone can discriminate between two mon-
However, the changes that take place dur- key faces (Pascalis, de Haan, & Nelson, 2002).
Summary ing the early years are particularly important However, 6-month-olds given 3 months of
In general, brain development begins a because they are the bedrock of what comes experience viewing a range of monkey faces
few weeks after conception and is thought to after. Higher level functions are dependent retain the recognition ability at 9 months
be complete by early adulthood. The basic on lower level functions, the evidence for (Pascalis et al., 2005). Thus, the plasticity
structure of the brain is laid down primarily which is primarily in the basic cognitive pro-
during the prenatal period and early child- cesses and sensory perceptual systems. When
hood, and the formation and refinement of infants are born, their brains are prepared for
neural networks continues over the long certain types of experience. For example, as
term. The brains’ many functions do not discussed below, infants’ brains are tuned to
develop at the same time nor do their devel- the sounds of virtually all languages, but with
opmental patterns follow the same time experience, their brains become most tuned
frame. Although basic sensation and per- to their native language (see Kuhl, 2004, for
ception systems are fully developed by the discussion). This perceptual bias is the basis
time children reach kindergarten age, other for learning language; the brain is partially
systems such as those involved in memory, tuned to be sensitive to language sounds but
decision making, and emotion continue to not so broadly tuned as to be sensitive to all
develop well into childhood. The founda- possible sounds.
tions of many of these abilities, however, are Subsequent language development builds
constructed during the early years. on this initial sensitivity. Within the first year
The principles of anatomical change of life, infants learn to discriminate among
described above are essential to the matu- sounds that are specific to the language they
Photo: © iStockphoto.com/Jelani Memory
ration and development of the brain. These are exposed to in their particular environ-
processes are in turn responsible for the ment. Before the time they are 6 months
development of a vast repertoire of behav- old, infants can discriminate among sounds
iors that characterizes the early years of life. of almost any language. Between 6 and 12
In terms of motor development, both synap- months, the brain begins to specialize in dis-
tic pruning and myelination are responsible criminating sounds of the native language
for the improved precision and speed of coor- and loses the ability to discriminate sounds
dinated movement. In addition, they are in nonnative languages (Kuhl, Tsao, & Liu,
important in the development of cognitive 2003). This narrowing of perceptual sensitiv-
skills. Improved perception of speech sounds ity is important because it is related to later
Early childhood represents a period par-
and face recognition, for example, are likely language ability in that better discrimination
ticularly important to development of a
the result of synaptic reorganization, a pro- of native language sounds predicts better lan-
healthy brain.
cess that is dependent on experience. guage skills later in life (Kuhl, 2004).
T
How the Timing and Quality of Early he effects of experience go beyond brain development that is in some way supe-
Experiences Influence the Development the simple modulation of plasticity. In rior. So although the BEIP studies do suggest
of Brain Architecture fact, experience shapes the structure that a lack of good quality experience is detri-
S. E. Fox, P. Levitt, & C. A. Nelson (in press). Child of the brain, a finding that has been demon- mental, they do not provide evidence for the
Development. strated by the Bucharest Early Intervention effects of enriched experience.
Project (BEIP). This ongoing longitudinal
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