The patterning of the neuroaxis (i.e.

, head to tail) is
for the most part completed by about the 5th prenatal
week.
 Proliferation
Once the neural tube has closed, cell division leads to a
massive proliferation of new neurons (neurogenesis),
generally beginning in the 5th prenatal week, and peaking
between the 3rd and 4th prenatal months
Proliferation
begins in the innermost portion of the neural
8 Neural Bases of Cognitive Development
tube, referred to as the ventricular zone (
 Neural development consists of seven stages [4]:

1. Cell birth (neurogenesis, gliogenesis)

2. Cell migration
3. Cell differentiation
4. Cell maturation (dendrite and axon growth)
5. Synaptogenesis (formation of synapses)
6. Cell death and synaptic pruning
7. Myelogenesis (formation of myelin)
 Schematic of projection neuron generation and migration in the mouse neocortex.

Kenneth Y. Kwan et al. Development 2012;139:1535-1546

2012.
 of the neural tube undergoes a dramatic transformation
specifically, three distinct vesicles are formed:
the proencephalon (forebrain), mesencephalon (midbrain),
and rhombencephalon (hindbrain).
proencephalon splitting into the
telencephalon, which will give rise to the cerebral hemispheres,
and the diencephalon, which gives rise to the
thalamus and hypothalamus. The rhombencephalon will
in turn give rise to the metencephalon (from which
the pons and cerebellum are derived) and the myelencephalon which will give rise to the
medulla). The mesencephalon
gives rise to the midbrain.
 Postnatal Neurogenesis
With the exception of cells in the olfactory bulb, it was
assumed until recently that the nervous system at birth
had virtually all the neurons it would ever have, and that
no new neurons were added

there is now widespread agreement that in certain

regions of the brain, new cells are added for many years
after birth. Where this agreement breaks down is in determining
precisely which regions experience this postnatal
birth of new neurons. There is little controversy
that the dentate gyrus of the hippocampus experiences
this process; controversy exists,
 Particularly relevant to this chapter is the observation
that the addition of such cells can be influenced by
experience (e.g., Gould et al., 1999; Mirescu, Peters, &
Gould, 2004).
2003). Collectively, experience
appears to have a profound influence on postnatal
neurogenesis.
 Cell Migration
The cortex proper (arguably the seat of cognition) is
formed by a process whereby newly formed cells migrate
out beyond their birthplace to ultimately give rise
to a six-layered cortex
At approximately
20 weeks gestation, the cortical plate consists of three
layers, and by the 7th prenatal month the final contingent
of six layers can be seen
There are two types of migratory patternsradial
and nonradial (generally tangential).
 The Growth and Development of Processes
Axons and Dendrites
Once a neuron has completed its migratory journey, it
generally proceeds along one of two roads: The cell can
differentiate and develop axons and dendrites (the topic
of this section), or it can be retracted through the normative
process of apoptosis (programmed cell death), a
phenomenon that is widespread (e.g., 40% to 60% of all
neurons may die
 The Development of Axons
Growth cones, small structures that sit on top of axons,
appear to play a key role both in developing the axon and
in guiding it to its target
The Development of Dendrites
Recent work has indicated that the gene calcium-regulated
transcriptional activator (CREST) plays an important
role in the development of dendrites

Dendritic sprouting begins to occur at approximately

15 weeks, about the same time that axons
reach the cortical plate.
 synaptic plasticity is driven by synaptic activity.
 reducing activity in a given neuron before
a synapse has been formed leads to a reduction in functional
inputs to that neuron (from neighboring neurons);
in contrast, reducing activity to a given neuron after a
synapse has been established leads to an increase in
synaptic input (Burrone, OByrne, & Murthy, 2002).
 Myelination
Myelin is a lipid/protein substance that wraps itself
around an axon as a form of insulation and, as a result,
increases conduction velocity
During the first postnatal year, regions of the brain stem
myelinate, as does the cerebellum and splenium of corpus
callosum; by 1 year, myelination of all regions of the
corpus callosum is underway.
 the assembly of the basic architecture of the
brain occurs during the first two trimesters of fetal life,
with the last trimester and the first few postnatal years
reserved for changes in connectivity and function. The
most prolonged changes occur in the wiring of the brain
(synaptogenesis) and in making the brain work more efficiently
(myelination), both of which show dramatic,
nonlinear changes from the preschool period through the
end of adolescence.