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feedback) reinforce the behavior and increase its likelihood of occurrence in the future. Bad consequences (negative feedback) have the opposite effect.Cockroaches learning to run through a simple maze to find food is a simple example of instrumental learning (also known as operant conditioning). Latent Learning involves memory of patterns or events when there is no apparent reward or punishment associated with the behavior. A sand wasp, for example, learns the location of her nest site by taking a short reconnaissance flight each time she leaves the nest. She remembers the pattern of surrounding landmarks to help her find the nest when she returns. Likewise, worker ants can remember a series of landmarks along a trail and follow them (in reverse order) back home to the nest site.Honey bees also show latent learning when they follow the waggle dance of a forager and then use that information to find the reported nectar source. Imprinting is a special case of programmed learning that occurs early in life and only within a short timewindow known as the critical period. During this brief interval, the animal acquires an indelible memory of certain salient stimuli in its home environment (taste of the host plant, smell of the nest site, etc.).This memory is retained throughout life and recalled later when needed. Fruit fly larvae, for example, will imprint on the taste and smell of their food.If reared on a diet that contains apple extract, adult females will show a strong preference for apples when they eventually search for a place to lay their own eggs.Not just any stimulus will do.Imprinting is apparently regulated by an innate neural template that restricts what can be remembered. Complex BehaviorEthologists are often careful to distinguish between learned and innate behaviors, but in reality the two are at extreme opposite ends of a single continuum. Most overt behavior is neither 100% innate nor 100% learned. Sometimes innate behaviors may be modified (or modulated) through practice and experience. In locusts, for example, the ability to fly is innate, but an older, experienced individual consumes less energy (per unit time) than a novice flier. This suggests that the older insect has learned to fly more efficiently. Similarly, learned behaviors may incorporate or depend upon elements of innate behavior. Indeed, the ability to learn, to associate, or to remember is almost certainly an innate feature of the insect39。s nervous system. Schematically, it may be useful to think of a box that represents the boundaries of an animal39。s ethogram. All behavior must occur inside the physiological limits of this box (. a beetle larva does not have wings, therefore it cannot fly). Within the box, a set of innate behaviors can be simplistically represented by straight lines. By following a zigzag route, an insect can use only innate behavior to get from point A to point B. But a learned behavior, superimposed on this innate grid, might provide a shortcut that is more useful or more efficient. As in the locust example above, the innate ability to fly may be refined and improved through experience