Learning also has costs that arise from the time, materials, and energy necessary to acquire and store information. 'Behavioral costs' arise from sampling a range of behaviors early in the learning process. For instance, birds take large amounts of time learning foraging skills and suffer energy losses from their mistakes; species that use a wider range of resources have more pronounced exploratory behavior during this period. Exploration may be costly in terms of time and energy, or in terms of increased risk of attack by natural enemies. 'Tissue costs' of learning arise from the neural tissue necessary to acquire and store information. Since neural tissue is particularly energetically expensive to maintain, tissue costs can be considerable.
Costs of learning translate into various tradeoffs with learning ability. In Drosophila fruit flies, artificial selection for faster learning in adults resulted in reduced competitive ability by larvae. In Pieris butterflies, natural variation in learning ability is causally linked with delays in reproduction.
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