Abstract

The ability of animals to explore landmarks in their environment is essential to their fitness. Landmarks are widely recognized to play a key role in navigation by providing information in multiple sensory modalities. However, what is a landmark? We propose that animals use a hierarchy of information based upon its utility and salience when an animal is in a given motivational state. Focusing on honeybees, we suggest that foragers choose landmarks based upon their relative uniqueness, conspicuousness, stability, and context. We also propose that it is useful to distinguish between landmarks that provide sensory input that changes (“near”) or does not change (“far”) as the receiver uses these landmarks to navigate. However, we recognize that this distinction occurs on a continuum and is not a clear-cut dichotomy. We review the rich literature on landmarks, focusing on recent studies that have illuminated our understanding of the kinds of information that bees use, how they use it, potential mechanisms, and future research directions.

Highlights

  • Sophisticated strategies and neural mechanisms have evolved to facilitate navigation. Depending on their natural history and ecology, many species need spatial memories to guide them on their paths

  • Recurring theme is that animals usually orient themselves towards their goals with multiple, redundant forms of information and prioritize this information according to its reliability and salience [1]

  • Our goal is to review the different senses in which the term landmark has been used and how landmarks and landscapes are used by honey bees

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Summary

What Are Landmarks?

A major and fascinating question in the study of animal cognition is how animals identify, remember, and use landmarks to navigate. Depending on their natural history and ecology, many species need spatial memories to guide them on their paths These navigational strategies use multiple sensory modalities, including vision, olfaction, proprioception, electroreception, and magnetoreception. The presence of these entities is used to help identify and define the location at which they are found [2,3] It is not known how animals perceive their goals relative to or nested within an overall landscape, but animals functionally use these entities at different stages along a path to reach a goal. Insects 2019, 10, 342 landscapes is not absolute and depending on the exact sensory stimuli, animals may perceive them as a continuum In many cases, it is unclear if a single stimulus aspect is enough to describe an entity or if entities consist of multiple stimuli that animals can recognize and use. We focus primarily on examples drawn from vision but have tried to define our overall concepts broadly to encompass multiple sensory modalities

How Have Landmarks Been Described?
Why Is It So Difficult to Define Landmarks?
A Proposed Definition for Landmarks
Representations of Space in Honeybees
Different Types of Navigation
Snapshots
Vectors
Features and the Sensory World of the Animal
Feature Hierarchies
Orientation Flights and Returning to Known Locations
Olfaction
Magnetoreception and Mechanosensation of Electrical Charges
Multimodality
Landmarks in Other Insects
Neural Bases of Landmark Memory
Recent Findings and Future Directions
Full Text
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