Cleanliness is next to godliness: mechanisms for staying clean.

Getting dirty is a fundamental problem, and one for which there are few solutions, especially across the enormous range of animal size. How do both a honeybee and a squirrel get clean? In this Review, we discuss two broad types of cleaning, considered from the viewpoint of energetics. Non-renewable cleaning strategies rely upon the organism as an energy source. Examples include grooming motions, wet-dog shaking or the secretion of chemicals. Renewable cleaning strategies depend on environmental sources of energy, such as the use of eyelashes to redirect incoming wind and so reduce deposition onto the eye. Both strategies take advantage of body hair to facilitate cleaning, and honeybees and squirrels, for example, each have around 3 million hairs. This hair mat increases the area on which particles can land by a factor of 100, but also suspends particles above the body, reducing their adhesion and facilitating removal. We hope that the strategies outlined here will inspire energy-efficient cleaning strategies in synthetic systems.