Path integration and optic flow in flying insects: a review of current evidence.

Path integration is a key navigation mechanism used by many animals, involving the integration of direction and distance of path segments to form a goal vector that allows an animal to return directly to its starting point. While well established for animals walking on solid ground, evidence for path integration in animals moving without ground contact, such as flying insects, is less clear. The review focuses on flying Hymenoptera, particularly bees, which are extensively studied. Although bees can use flight distance and direction information, evidence for genuine path integration is limited. Accurately assessing distance travelled is a major challenge for flying animals, because it relies on optic flow-the movement of visual patterns across the eye caused by locomotion. Optic flow depends on both the animal's speed and the spatial layout of the environment, making it ambiguous for precise distance measurement. While path integration is crucial for animals like desert ants navigating sparse environments with few navigational cues, we argue that flying Hymenopterans in visually complex environments, rich in objects and textures, rely on additional navigational cues rather than precise path integration. As they become more familiar with an environment, they may iteratively refine unreliable distance estimates derived from optic flow. By combining this refined information with directional cues, they could determine a goal vector and improve their ability to navigate efficiently between key locations. In the case of honeybees, this ability also enables them to communicate these refined goal vectors to other bees through the waggle dance.