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. 2014 Apr 3;8:97. doi: 10.3389/fncel.2014.00097

Table 1.

Effects of environmental enrichment on neurobiology and behavior.

Study’s primary objective Animal species/strain EE methods Frequency of changing EE method Protein/behavioral parameters investigated Significant findings Reference
• Effects of EE on neurodegeneration during AD • AD11 mice • Large cages with wire mesh lid, several food hoppers, running wheel and objects of different shapes (tunnels, shelters, stairs, boxes) • Once per week • Visual object recognition test • ↑ Visual object recognition memory and spatial memory Berardi et al. (2007)
• Morris water maze test • ↓ Aβ deposition in hippocampus
• IHC • ↓ Progression of neurodegeneration
• Effects of long-term EE on hippocampal neurogenesis • 10 months old female C57BL/6 mice • Large cages, with re-arrangeable set of plastic tubes, a running wheel, nesting material, and toys • Behavioral testing with Activity chamber, Rotarod, and Water maze • Fivefold increase in hippocampal neurogenesis in enriched environment Kempermann et al. (2002)
• IHC and IF for lipofuscin deposits in neurons • ↑ Learning, exploratory behavior, and locomotor activity
• ↓ Lipofuscin deposits in the dentate gyrus
• Effects of environmental complexity on spatial abilities, dendritic arborization, and spine density • Wistar rat (21 days old) • Ten rats in a large cage of two levels connected by ramps, containing wood shavings, a running wheel, a shelter, plastic colored toys, and constructions • Once a week • Full-baited maze procedure and forced-choice procedure, performed in a radial maze • ↑ Performance in the Radial maze and Morris water maze tasks Leggio et al. (2005)
• Morris water maze • ↑ Dendritic arborization and spine density in layer-III parietal pyramidal neurons
In vivo Golgi-like filling of the neurons for the visualization of dendritic arborization
• Influence of EE on neurotrophins levels in the cerebellum • Adult male Wistar rats • 10 rats in a large cage with two levels, connected by ramps, contain wood shavings, a running wheel, a shelter, plastic colored toys, and constructions • Twice per week • Determination of BDNF and NGF in all brain regions using ELISA • ↑ BDNF levels in the cerebellum, frontal cortex and hippocampus Angelucci et al. (2009)
• ↓ BDNF levels in the striatum
• ↑ NGF levels in the cerebellum and striatum
• No significant change in NGF levels in the frontal cortex and hippocampus
• Effects of EE on the neurogenesis and the extracellular concentrations of glutamate and GABA in the hippocampus • Male Wistar rats of 2 and 25 months • Two running wheels, a re-arrangeable set of plastic tunnels, an elevated platform, toys • Every 3–4 days • Water maze test • ↑ Spatial memory performance Segovia et al. (2006)
• Neurogenesis in the dentate gyrus of hippocampus using BrdU labeling • ↑ Hippocampal neurogenesis in both young and aged enriched rats
• Glutamate and GABA concentration in CA3 region of hippocampus using microdialysis probes • ↓ Hippocampal neurogenesis in control rats
• No effect of EE on basal concentration of Glutamate and GABA in young rats
• ↑ Basal glutamate and GABA concentration in old rats
• Effects of ageing and EE on synaptic plasticity • Male Fischer rats • Large cages furnished with various toys and small constructions • Electron microscopic morphometry for the analyses of density and sizes of synapses • ↓ Synaptic vesicle density with age Nakamura et al. (1999)
• EE restored age-related loss of synaptophysin

EE, environmental enrichment; AD, Alzheimer’s disease; IHC, immunohistochemistry; Aβ, amyloid-β; IF, immunofluorescence; BDNF, rain-derived neurotrophic factor; NGF, nerve growth factor; ELISA, enzyme linked immunosorbent assay; BrdU, bromodeoxyuridine; GABA, gamma-amino butyric acid; CA, Cornu Ammonis.

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