The tongue as a gripper

Frogs, chameleons and anteaters are striking examples of animals that can grab food using only their tongue. How does the soft and wet surface of a tongue grip onto objects before they are ingested? Here, we review the diversity of tongue projection methods, tongue roughnesses and tongue coatings, our goal being to highlight conditions for effective grip and mobility. A softer tongue can reach farther: the frog Rana pipiens tongue is 10 times softer than the human tongue and can extend to 130% of its length when propelled in a whip-like motion. Roughness can improve a tongue's grip: the spikes on a penguin Eudyptes chrysolophus tongue can be as large as fingernails, and help the penguin swallow fish. The saliva coating on the tongue, a non-Newtonian biofluid, can either lubricate or adhere to food. Frog saliva is 175 times more viscous than human saliva, adhering the tongue to slippery, furry or feathery food. We pay particular attention to using mathematical models such as the theory of capillarity, elasticity and friction to elucidate the parameters for effective tongue use across a variety of vertebrate species. Finally, we postulate how the use of wet and rough surfaces to simultaneously sense and grip may inspire new strategies in emerging technologies such as soft robots.