of functioning of the brain. Using astrocytes (cells that support neuron functioning by providing nutrients and removing waste) as the index, there are higher amounts of astrocyte per neuron in the complex-environment animals than in the caged groups. Overall, these studies depict an orchestrated pattern of increased capacity in the brain that depends on experience.

Other studies of animals show other changes in the brain through learning; see Box 5.1. The weight and thickness of the cerebral cortex can be measurably altered in rats that are reared from weaning, or placed as adults, in a large cage enriched by the presence both of a changing set of objects for play and exploration and of other rats to induce play and exploration (Rosenzweig and Bennett, 1978). These animals also perform better on a variety of problem-solving tasks than rats reared in standard laboratory cages. Interestingly, both the interactive presence of a social group and direct physical contact with the environment are important factors: animals placed in the enriched environment alone showed relatively little benefit; neither did animals placed in small cages within the larger environment (Ferchmin et al., 1978; Rosenzweig and Bennett, 1972). Thus, the gross structure of the cerebral cortex was altered both by exposure to opportunities for learning and by learning in a social context.

Does Mere Neural Activity Change the Brain or Is Learning Required?

Are the changes in the brain due to actual learning or to variations in aggregate levels of neural activity? Animals in a complex environment not only learn from experiences, but they also run, play, and exercise, which activates the brain. The question is whether activation alone can produce brain changes without the subjects actually learning anything, just as activation of muscles by exercise can cause them to grow. To answer this question, a group of animals that learned challenging motor skills but had relatively little brain activity was compared with groups that had high levels of brain activity but did relatively little learning (Black et al., 1990). There were four groups in all. One group of rats was taught to traverse an elevated obstacle course; these “acrobats” became very good at the task over a month or so of practice. A second group of “mandatory exercisers” was put on a treadmill once a day, where they ran for 30 minutes, rested for 10 minutes, then ran another 30 minutes. A third group of “voluntary exercisers” had free access to an activity wheel attached directly to their cage, which they used often. A control group of “cage potato” rats had no exercise.

What happened to the volume of blood vessels and number of synapses per neuron in the rats? Both the mandatory exercisers and the voluntary exercisers showed higher densities of blood vessels than either the cage potato rats or the acrobats, who learned skills that did not involve significant



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