Advances in Bioscience and Biotechnology, 2013, 4, 785-797 ABB
doi:10.4236/abb.2013.47103 Published Online July 2013 (http://www.scirp.org/journal/abb/ )
Selective and constructive mechanisms contribute to neural
circuit formation in the barrel cortex of the developing rat
Eileen Uribe-Querol
1*
, Eduardo Martínez-Martínez
2
, Luis Rodrigo Hernández
2
,
Patricia Padilla Cortés
3
, Horacio Merchant-Larios
2
, Gabriel Gutiérrez-Ospina
2
1
División de Estudios de Posgrado e Investigación, Facultad de Odontología, Universidad Nacional Autónoma de México, Mexico
City, Mexico
2
Cell Biology and Physiology Department, Instituto de Investigaciones Biomédicas, Universidad Nacional Autónoma de México,
Mexico City, Mexico
3
Unidad de HPLC, Instituto de Investigaciones Biomédicas, Universidad Nacional Autónoma de México, Mexico City, Mexico
Email:
*
euquerol@comunidad.unam.mx
Received 26 May 2013; revised 28 June 2013; accepted 7 July 2013
Copyright © 2013 Eileen Uribe-Querol et al. This is an open access article distributed under the Creative Commons Attribution Li-
cense, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.
ABSTRACT
The cellular strategy leading to formation of neuronal
circuits in the rodent barrel cortex is still a matter of
controversy. Both selective and constructive mecha-
nisms have been proposed. The selective mechanism
involves an overproduction of neuronal processes and
synapses followed by activity dependent pruning.
Conversely, a constructive mechanism would increase
the number of axons, dendrites, and synapses during
development to match functionality. In order to dis-
cern the contributions of these two mechanisms in
establishing a neuronal circuit in the somatosensory
cortex, morphometric analysis of dendritic and ax-
onal arbor growth was performed. Also, the number
of synapses was followed by electron microscopy
during the first month of life. We observed that ax-
onal and dendritic arbors retracted distal branches,
and elongated proximal branches, resulting in in-
creased arbor complexity. This neuronal remodeling
was accompanied by the steady increase in the num-
ber of synapses within barrel hollows. Similarly, the
content of molecular markers for dendrites, axons
and synapses also increased during this period. Fi-
nally, cytochrome oxidase activity rose with age in
barrels indicating that the arbors became more com-
plex while synapse density and metabolic demands
increased. Our results support the simultaneous use
of both selective and constructive mechanisms in es-
tablishing the barrel cortex circuitry.
Keywords: Barrel Cortex; Synaptogenesis; Axonal
Growth; Dendritic Growth; Thalamocortical Pathway
1. INTRODUCTION
During the ontogeny of the nervous system, axonal and
dendritic branches, as well as synapses, are thought to be
formed in numbers that far exceed the amount required
by the mature brain to function properly. The predomi-
nant theory for neuronal circuit formation is the selective
mechanism, which states that a period of overproduction
of neuronal processes and synapses is followed by the
elimination of a significant fraction of them. This process
of elimination is driven by competition. Axons or den-
drites compete for synaptic partners that may provide
neurotrophic factors in sufficient amount to consolidate
synapses and retain their partners [1]. Axonal and den-
dritic branches that establish synaptic-like contacts with
incorrect partners are eliminated because they do not re-
ceive enough trophic support. Even though the notion
described above constitutes the dominant view about
how the developing brain sculpts its circuitry, increasing
anatomical evidence supports that the selective mecha-
nism is not a universal rule. Indeed, anatomical evidence
gathered during the development of different brain struc-
tures indicates that axons may grow towards and within
their targets by preferentially adding branches with a
high degree of precision until they attain their adult size
[2]. Similarly, multiple reports also support the notion
that the dendritic arbors of some classes of neurons also
develop preferentially by adding individual branches
until they reach their adult configuration [3]. The rodent
primary somatosensory cortex (S1) processes tactile in-
formation and contains a representation of the body
formed by modules called barrels. Each barrel contains
an anatomical map that represents collections of mech-
anosensory receptors distributed throughout the body. *
Corresponding author.
Published Online July 2013 in SciRes. http://www.scirp.org/journal/abb