Table 1.
Studies on environmental enrichment’s effects in healthy animals.
| Reference | Species (Age or Weight at the Start of Environmental Enrichment) |
Environmental Enrichment Type (Duration) |
Environmental Enrichment Effects on Cerebellar Structure and Function (Age at the Effect Evaluations) |
|---|---|---|---|
| Uphouse, 1978 [68] | Male Fischer rats (28 days) |
Environmental enrichment —without running wheels; with novelty manipulation (32 days) |
Increased chromatin level (2 months) |
| Uphouse and Tedeschi, 1979 [69] | Male Fischer rats (28 days) |
Environmental enrichment —without running wheels; with novelty manipulation (60 days) |
Unchanged chromatin level (about 3 months) |
| Naka et al., 2002 [67] | Male ICR mice (28 days) |
Environmental enrichment —with running wheels and novelty manipulation (40 days) |
Increased noradrenaline expression; unchanged serotonin and metabolites expression (about 2 months) |
| Nithianantharajah et al., 2004 [64] | Female C57BL/6 mice (28 days) |
Environmental enrichment —with running wheels and novelty manipulation (30 days) |
Unchanged synaptophysin level (about 2 months) |
| Angelucci et al., 2009 [57] | Male Wistar rats (21 days) |
Environmental enrichment —with running wheels and novelty manipulation (about 120 days) |
Increased nerve growth factor (NGF) and brain-derived neurotrophic factor (BDNF) levels (140 days) |
| Pascual and Bustamante, 2013 [65] | Male Sprague–Dawley rats (22 days) |
Environmental enrichment —with running wheels and novelty manipulation (10 days) |
Unchanged anxiety-like behavior (33–34 days); unchanged vermal Purkinje cell dendritic outgrowth (36 days) |
| Vazquez-Sanroman et al., 2013 [66] | Male Balb/c AnNHsd mice (21 days) |
Environmental enrichment —without running wheels; with novelty manipulation (1/4/8 weeks) |
After 1 week: unchanged pro-BDNF and mature BDNF proteins; increased BDNF immunoreactivity at granular layer; unchanged BDNF immunoreactivity at Purkinje layer (4 weeks) After 4 weeks: increased BDNF immunoreactivity at granular and Purkinje layers (7 weeks) After 8 weeks: increased pro-BDNF and mature BDNF proteins; increased BDNF immunoreactivity at granular and Purkinje layers (11 weeks) |
| De Bartolo et al., 2015 [62] | Male Wistar rats (21 days) |
Environmental enrichment —with running wheels and novelty manipulation (about 100 days) |
Increased cerebellar Purkinje cell dendritic spine density and size (about 120 days) |
| Horvat et al., 2015 [72] | Male Wistar rats (6 months) |
Environmental enrichment —without running wheels; with novelty manipulation (21 days) |
Increased pituitary adenylate cyclase activating polypeptide (PACAP) 27 expression; unchanged PACAP 38 expression (27 weeks) |
| Scholz et al., 2015 [71] | Male C57BL/B6 mice (7 weeks) |
Environmental enrichment —with running wheels and novelty manipulation—based on a three-level maze, without objects (24 h; 21 days) |
Improved spatial learning (10 weeks); decreased volume (about 7 weeks; 10 weeks) |
| Eshra et al., 2019 [70] | C57BL/6 mice (at birth) |
Environmental enrichment —with running wheels and novelty manipulation (70–80 days) |
Improved motor performance; higher granule cell firing frequency (70–80 days) |
| Kim et al., 2019 [63] | C57BL/6 mice (3 weeks) |
Environmental enrichment —with running wheels and novelty manipulation (28 days) |
Selective increase in parallel fiber-to-Purkinje cell synapses of same dendritic origin, with local synaptic strengthening (7 weeks) |
Note: unless otherwise specified, the described effects involve the entire cerebellar structure.