SHORT-TERM EXPOSURE TO AN ENRICHED ENVIRONMENT
ENHANCES DENDRITIC BRANCHING BUT NOT BRAIN-DERIVED
NEUROTROPHIC FACTOR EXPRESSION IN THE HIPPOCAMPUS
OF RATS WITH VENTRAL SUBICULAR LESIONS
B. BINDU, P. A. ALLADI, B. M. MANSOORALIKHAN,
B. N. SRIKUMAR, T. R. RAJU AND B. M. KUTTY*
Department of Neurophysiology, National Institute of Mental Health
and Neuro Sciences, P.O. Box 2900, Hosur Road, Bangalore 560 029,
India
Abstract—Environmental enrichment promotes structural
and behavioral plasticity in the adult brain. We have evalu-
ated the efficacy of enriched environment on the dendritic
morphology and brain-derived neurotrophic factor (BDNF)
expression in the hippocampus of ventral subicular–lesioned
rats.
Bilateral ventral subicular lesion has significantly reduced
the dendritic architecture and spine density of hippocampal
pyramidal neurons. The lesioned rats exposed to enriched
housing for 10 days showed a significant degree of morpho-
logical plasticity in terms of enhanced dendritic branching
and spine density. However, the BDNF expression in the
hippocampus remained unchanged following subicular le-
sion and following environmental enrichment. We suggest
the participation of other neurotrophic factors in mediating
the synaptic plasticity events following exposure to environ-
mental enrichment in ventral subicular–lesioned rats. © 2006
IBRO. Published by Elsevier Ltd. All rights reserved.
Key words: brain lesion, dendritic atrophy, neuronal plasticity.
Environmental enrichment has been extensively used to
demonstrate the behavioral and brain plasticity in response to
experience (Rosenzweig et al., 1962; Greenough et al.,
1973). Various studies have reported that adult rats exposed
to a complex environment consisting of a combination of
social stimulation and physical activity induce chemical and
anatomical changes and enhanced behavioral perfor-
mance (Connor and Diamond, 1982; Faherty et al., 2003;
Fiala et al., 1978; Knafo et al., 2001; Rampon et al., 2000;
Bennett et al., 1969; Diamond et al., 1964, 1966, 1975;
Altman and Das, 1965). Additionally different types of en-
richment may have different effects as reported by Bennett
et al. (2006). They have observed an overall enhancement
of spatial memory functions in aged mice when exposed
continuously for a long period to enriched environment
(EE) than providing daily enrichment. The neuroanatomi-
cal changes are attributed to alterations in gene expres-
sion linked to neuronal structure, synaptic plasticity and
transmission (Rampon and Tsien, 2000). Changes in neu-
rotrophins’ mRNA levels in the cortex and hippocampus
may also play an important role in experience induced
shaping of neuronal connection (Torasdotter et al., 1996).
The neurotrophins like brain-derived neurotrophic factor
(BDNF) and nerve growth factor play a major role in syn-
aptic plasticity in the hippocampus and other brain regions
such as somatosensory and visual cortex (McAllister et al.,
1995, 1996).
Experience-associated changes in brain morphology
leading to functional recovery have been reported after
brain injury (Kolb and Gibb, 1991; Kolb et al., 1998;
Passineau et al., 2001). Rats subjected to environmental
enrichment following transient global cerebral ischemia
exhibited enhanced morphological and behavioral plastic-
ity (Briones et al., 2000). Enriched rehabilitation following
middle cerebral artery occlusion improved motor perfor-
mances in staircase reaching and beam traversing tasks in
ischemic rats. Enriched housing has in fact produced en-
hanced dendritic complexity in the layer V pyramidal cells
within undamaged motor cortex. Task specific rehabilita-
tion augments neuronal plasticity in the affected region of
the brain and thereby promotes functional outcome (Bier-
naskie and Corbett, 2001). Similarly, studies have reported
increased dendritic complexity in the undamaged sensori-
motor cortex following exposure to EE in rats with cortical
lesion and cerebral infarction (Jones and Schallert, 1994;
Johansson, 1996). Young et al. (1999) have also reported
the various plasticity evoking properties of environmental
enrichment following brain insults. They have suggested
that enriched housing helps to optimize the structural mor-
phology in the intact brain. Gobbo and O’Mara (2005)
found that EE enhances functional recovery after ischemia
and they attributed the recovery to enhanced BDNF ex-
pression following environmental enrichment. BNDF/ty-
rosine kinase B signaling has been shown to mediate
morphological plasticity in the hippocampal neurons (Tyler
and Pozzo-Miller, 2001). Additionally this may act as an
environmental stimuli to preserve some aspects of molec-
ular machinery responsible for neuronal plasticity (Restivo
et al., 2005).
*Corresponding author. Tel: +91-80-26995170; fax: +91-80-2656-
2121 or +91-80-2656-4830.
E-mail address: bindu@nimhans.kar.nic.in (B. M. Kutty).
Abbreviations: BDNF, brain-derived neurotrophic factor; CA1, cornu
ammonis 1; CA3, cornu ammonis 3; EE, enriched environment; FITC,
fluorescein thiocyanate; NC, normal control; PBS, phosphate-buffered
saline; PBSTx, phosphate-buffered saline with Triton X-100; TBS–
Tween, 250 nM Tris–HCl, 0.9% NaCl and 0.1% Tween 20; VC, vehicle
control; VSL, ventral subicular–lesioned; VSL+EE, ventral subicular–
lesioned rats reared in enriched housing condition; VSL+SH, ventral
subicular–lesioned rats reared in standard housing condition.
Neuroscience 144 (2007) 412– 423
0306-4522/07$30.00+0.00 © 2006 IBRO. Published by Elsevier Ltd. All rights reserved.
doi:10.1016/j.neuroscience.2006.09.057
412