Skin wrinkles and folds enable asymmetric stretch in the elephant trunk.

The elephant's trunk is multifunctional: It must be flexible to wrap around vegetation, but tough to knock down trees and resist attack. How can one appendage satisfy both constraints? In this combined experimental and theoretical study, we challenged African elephants to reach far-away objects with only horizontal extensions of their trunk. Surprisingly, the trunk does not extend uniformly, but instead exhibits a dorsal "joint" that stretches 15% more than the corresponding ventral section. Using material testing with the skin of a deceased elephant, we show that the asymmetry is due in part to patterns of the skin. The dorsal skin is folded and 15% more pliable than the wrinkled ventral skin. Skin folds protect the dorsal section and stretch to facilitate downward wrapping, the most common gripping style when picking up items. The elephant's skin is also sufficiently stiff to influence its mechanics: At the joint, the skin requires 13 times more energy to stretch than the corresponding length of muscle. The use of wrinkles and folds to modulate stiffness may provide a valuable concept for both biology and soft robotics.

Jumping of flea beetles onto inclined platforms.

The flea beetle, Altica cirsicola, escapes predators by jumping and landing in a dense maze of leaves. How do they land on such varied surfaces? In this experimental study, we filmed the take-off, flight, and landing of flea beetles on a configurable angled platform. We report three in-flight behaviors: winged, wingless, and an intermediate winged mode. These modes significantly affected take-off speed, acceleration, and the duration that wings were deployed. When wings were closed, flea beetles rolled or pitched up to five times in the air. This work may help to understand how insects can jump and right themselves onto variable surfaces.

Drying dynamics of pellet feces.

Pellet feces are generated by a number of animals important to science or agriculture, including mice, rats, goats, and wombats. Understanding the factors that lead to fecal shape may provide a better understanding of animal health and diet. In this combined experimental and theoretical study, we test the hypothesis that pellet feces are formed by drying processes in the intestine. Inspirational to our work is the formation of hexagonal columnar jointings in cooling lava beds, in which the width L of the hexagon scales as L ∼ J-1 where J is the heat flux from the bed. Across 22 species of mammals, we report a transition from cylindrical to pellet feces if fecal water content drops below 0.65. Using a mathematical model that accounts for water intake rate and intestinal dimensions, we show pellet feces length L scales as L ∼ J-2.08 where J is the flux of water absorbed by the intestines. We build a mimic of the mammalian intestine using a corn starch cake drying in an open trough, finding that corn starch pellet length scales with water flux-0.46. The range of exponents does not permit us to conclude that formation of columnar jointings is similar to the formation of pellet feces. Nevertheless, the methods and physical picture shown here may be of use to physicians and veterinarians interested in using feces length as a marker of intestinal health.

Urinary flow through urethras with a rough lumen.

AIMS: This study investigates how lumen roughness and urethral length influence urinary flow speed. METHODS: We used micro-computed tomography scans to measure the lumen roughness and dimensions for rabbits, cats, and pigs. We designed and fabricated three-dimensional-printed urethra mimics of varying roughness and length to perform flow experiments. We also developed a corresponding mathematical model to rationalize the observed flow speed. RESULTS: We update the previously reported relationship between body mass and urethra length and diameter, now including 41 measurements for urethra length and 10 measurements for diameter. We report the relationship between lumen diameter and roughness as a function of position down the urethra for rabbits, cats, and pigs. The time course of urinary speed from our mimics is reported, as well as the average speed as a function of urethra length. CONCLUSIONS: Based on the behavior of our mimics, we conclude that the lumen roughness in mammals reduces flow speed by up to 25% compared to smooth urethras. Urine flows fastest when the urethra length exceeds 25 times its diameter. Longer urethras do not drain faster due to viscous effects counteracting the additional gravitational head. However, flows with our urethra mimics are still 6 times faster than those observed in nature, suggesting that further work is needed to understand flow resistance in the urethra.

Intestines of non-uniform stiffness mold the corners of wombat feces.

The bare-nosed wombat (Vombatus ursinus) is a fossorial, herbivorous, Australian marsupial, renowned for its cubic feces. However, the ability of the wombat's soft intestine to sculpt flat faces and sharp corners in feces is poorly understood. In this combined experimental and numerical study, we show one mechanism for the formation of corners in a highly damped environment. Wombat dissections show that cubes are formed within the last 17 percent of the intestine. Using histology and tensile testing, we discover that the cross-section of the intestine exhibits regions with a two-fold increase in thickness and a four-fold increase in stiffness, which we hypothesize facilitates the formation of corners by contractions of the intestine. Using a mathematical model, we simulate a series of azimuthal contractions of a damped elastic ring composed of alternating stiff and soft regions. Increased stiffness ratio and higher Reynolds number yield shapes that are more square. The corners arise from faster contraction in the stiff regions and relatively slower movement in the center of the soft regions. These results may have applications in manufacturing, clinical pathology, and digestive health.

Cats use hollow papillae to wick saliva into fur.

The cat tongue is covered in sharp, rear-facing spines called papillae, the precise function of which is a mystery. In this combined experimental and theoretical study, we use high-speed film, grooming force measurements, and computed tomography (CT) scanning to elucidate the mechanism by which papillae are used to groom fur. We examine the tongues of six species of cats from domestic cat to lion, spanning 30-fold in body weight. The papillae of these cats each feature a hollow cavity at the tip that spontaneously wicks saliva from the mouth and then releases it onto hairs. The unique shape of the cat's papillae may inspire ways to clean complex hairy surfaces. We demonstrate one such application with the tongue-inspired grooming (TIGR) brush, which incorporates 3D-printed cat papillae into a silicone substrate. The TIGR brush experiences lower grooming forces than a normal hairbrush and is easier to clean.

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.

Mammals repel mosquitoes with their tails.

The swinging of a mammal's tail has long been thought to deter biting insects, which, in cows, can drain up to 0.3 liters of blood per day. How effective is a mammal's tail at repelling insects? In this combined experimental and theoretical study, we filmed horses, zebras, elephants, giraffes and dogs swinging their tails. The tail swings at triple the frequency of a gravity-driven pendulum, and requires 27 times more power input. Tails can also be used like a whip to directly strike at insects. This whip-like effect requires substantial torques from the base of the tail on the order of 101-102 N m, comparable to the torque of a sedan, but still within the physical limits of the mammal. Based on our findings, we designed and built a mammal tail simulator to simulate the swinging of the tail. The simulator generates mild breezes of 1 m s-1, comparable to a mosquito's flight speed, and sufficient to deter up to 50% of mosquitoes from landing. This study may help us determine new mosquito-repelling strategies that do not depend on chemicals.

Chewing Holes for Camouflage.

Camouflaged objects are harder to detect if the background itself is more heterogeneous, and search becomes increasingly inefficient when the scene contains multiple items resembling the target. Some adult leaf beetles (Coleoptera: Chrysomelidae) with highly specialized habits make holes on host plant leaves while feeding. We propose that leaf beetles camouflage themselves with their feeding holes. The presence of holes makes predators' visual search harder, thus giving beetles more time to escape from the leaf surface either by jumping (Galerucinae: Alticini) or rolling (rest of Chrysomelidae). Based on behavioral observations and analysis of 25 photographs of feeding leaf beetles (15 species), we demonstrate that adult leaf beetles camouflage themselves by creating holes of uniform size, approximately half of the beetle body size. Observation of the feeding behavior and anatomy of a typical hole-feeding beetle (Altica cirsicola) showed that the foregut volume and head-prothorax mobility of beetles are the two major factors that constrain the hole size. A computer-simulated visual search test showed that the greater the number of holes, and the more each hole approached beetle body size, the longer it took humans (as models) to locate a beetle on a leaf. This study reports a newly discovered kind of camouflage, hole-feeding camouflage, in leaf beetles, which makes visual detection or recognition more difficult by changing the environmental background. This type of camouflage may open up a range of new possibilities for studies in animal cognition analysis and evolution of anti-predation defenses.

Modulation of orthogonal body waves enables high maneuverability in sidewinding locomotion.

Many organisms move using traveling waves of body undulation, and most work has focused on single-plane undulations in fluids. Less attention has been paid to multiplane undulations, which are particularly important in terrestrial environments where vertical undulations can regulate substrate contact. A seemingly complex mode of snake locomotion, sidewinding, can be described by the superposition of two waves: horizontal and vertical body waves with a phase difference of ± 90°. We demonstrate that the high maneuverability displayed by sidewinder rattlesnakes (Crotalus cerastes) emerges from the animal's ability to independently modulate these waves. Sidewinder rattlesnakes used two distinct turning methods, which we term differential turning (26° change in orientation per wave cycle) and reversal turning (89°). Observations of the snakes suggested that during differential turning the animals imposed an amplitude modulation in the horizontal wave whereas in reversal turning they shifted the phase of the vertical wave by 180°. We tested these mechanisms using a multimodule snake robot as a physical model, successfully generating differential and reversal turning with performance comparable to that of the organisms. Further manipulations of the two-wave system revealed a third turning mode, frequency turning, not observed in biological snakes, which produced large (127°) in-place turns. The two-wave system thus functions as a template (a targeted motor pattern) that enables complex behaviors in a high-degree-of-freedom system to emerge from relatively simple modulations to a basic pattern. Our study reveals the utility of templates in understanding the control of biological movement as well as in developing control schemes for limbless robots.

Hydrodynamics of defecation.

Animals discharge feces within a range of sizes and shapes. Such variation has long been used to track animals as well as to diagnose illnesses in both humans and animals. However, the physics by which feces are discharged remain poorly understood. In this combined experimental and theoretical study, we investigate the defecation of mammals from cats to elephants using the dimensions of large intestines and feces, videography at Zoo Atlanta, cone-on-plate rheological measurements of feces and mucus, and a mathematical model of defecation. The diameter of feces is comparable to that of the rectum, but the length is double that of the rectum, indicating that not only the rectum but also the colon is a storage facility for feces. Despite the length of rectum ranging from 4 to 40 cm, mammals from cats to elephants defecate within a nearly constant duration of 12 ± 7 seconds (N = 23). We rationalize this surprising trend by our mathematical model, which shows that feces slide along the large intestine by a layer of mucus, similar to a sled sliding down a chute. Larger animals have not only more feces but also thicker mucus layers, which facilitate their ejection. Our model accounts for the shorter and longer defecation times associated with diarrhea and constipation, respectively. This study may support clinicians use of non-invasive procedures such as defecation time in the diagnoses of ailments of the digestive system.

Duration of urination does not change with body size.

Many urological studies rely on models of animals, such as rats and pigs, but their relation to the human urinary system is poorly understood. Here, we elucidate the hydrodynamics of urination across five orders of magnitude in body mass. Using high-speed videography and flow-rate measurement obtained at Zoo Atlanta, we discover that all mammals above 3 kg in weight empty their bladders over nearly constant duration of 21 ± 13 s. This feat is possible, because larger animals have longer urethras and thus, higher gravitational force and higher flow speed. Smaller mammals are challenged during urination by high viscous and capillary forces that limit their urine to single drops. Our findings reveal that the urethra is a flow-enhancing device, enabling the urinary system to be scaled up by a factor of 3,600 in volume without compromising its function. This study may help to diagnose urinary problems in animals as well as inspire the design of scalable hydrodynamic systems based on those in nature.

Ant aggregations self-heal to compensate for the Ringelmann effect.

Fire ants, Solenopsis invicta, link their bodies together to form structures such as rafts, bivouacs and bridges. Such structures are in danger of being damaged by natural disturbances such as passing water currents. In this combined experimental and theoretical study, we investigate the self-healing of ant assemblages. We press two ant aggregations together and measure the forces to pull them apart. As the group size increases, the contribution of each ant decreases. This phenomenon, known as the Ringelmann effect, or social loafing, has previously been shown for cattle and humans. In this study, we show that it is a challenge for ants as well. We rationalize this effect with an agent-based simulation which exhibits the Ringelmann effect of ants that periodically make and break links with each other, but grip with higher probability if the ants are stretched. Over time, ants compensate for the Ringelmann effect by building more links. We use a mathematical model to show that the rate of new links is proportional to the number of free ants in the cluster. The principles found here may inspire new directions in self-healing and active materials.

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.

Mosquitoes survive raindrop collisions by virtue of their low mass.

In the study of insect flight, adaptations to complex flight conditions such as wind and rain are poorly understood. Mosquitoes thrive in areas of high humidity and rainfall, in which raindrops can weigh more than 50 times a mosquito. In this combined experimental and theoretical study, we here show that free-flying mosquitoes can survive the high-speed impact of falling raindrops. High-speed videography of those impacts reveals a mechanism for survival: A mosquito's strong exoskeleton and low mass renders it impervious to falling drops. The mosquito's low mass causes raindrops to lose little momentum upon impact and so impart correspondingly low forces to the mosquitoes. Our findings demonstrate that small fliers are robust to in-flight perturbations.

Fire ants actively control spacing and orientation within self-assemblages.

To overcome obstacles and survive harsh environments, fire ants link their bodies together to form self-assemblages such as rafts, bridges and bivouacs. Such structures are examples of self-assembling and self-healing materials, as ants can quickly create and break links with one another in response to changes in their environment. Because ants are opaque, the arrangement of the ants within these three-dimensional networks was previously unknown. In this experimental study, we applied micro-scale computed tomography, or micro-CT, to visualize the connectivity, arrangement and orientation of ants within an assemblage. We identified active and geometric mechanisms that ants use to obtain favorable packing properties with respect to well-studied packing of inert objects such as cylinders. Ants use their legs to push against their neighbors, doubling their spacing relative to random packing of cylinders. These legs also permit active control of their orientation, an ability ants use to arrange themselves perpendicularly rather than in parallel. Lastly, we found an important role of ant polymorphism in promoting self-aggregation: a large distribution of ant sizes permits small ants to fit between the legs of larger ants, a phenomenon that increases the number of average connections per ant. These combined mechanisms lead to low packing fraction and high connectivity, which increase raft buoyancy and strength during flash floods.

Fire ants self-assemble into waterproof rafts to survive floods.

Why does a single fire ant Solenopsis invicta struggle in water, whereas a group can float effortlessly for days? We use time-lapse photography to investigate how fire ants S. invicta link their bodies together to build waterproof rafts. Although water repellency in nature has been previously viewed as a static material property of plant leaves and insect cuticles, we here demonstrate a self-assembled hydrophobic surface. We find that ants can considerably enhance their water repellency by linking their bodies together, a process analogous to the weaving of a waterproof fabric. We present a model for the rate of raft construction based on observations of ant trajectories atop the raft. Central to the construction process is the trapping of ants at the raft edge by their neighbors, suggesting that some "cooperative" behaviors may rely upon coercion.

Onggi's permeability to carbon dioxide accelerates kimchi fermentation.

Since ancient times, Korean chefs have fermented foods in an onggi, a traditional earthenware vessel. The porous structure of the onggi mimics the loose soil where lactic acid bacteria is naturally found. This permeability has been purported to facilitate the growth of lactic acid bacteria, but the details of the process remain poorly understood. In this combined experimental and theoretical study, we ferment salted napa cabbage in onggi and hermetic glassware and measure the time course of carbon dioxide concentration, which is a signature of fermentation. We present a mathematical model for carbon dioxide generation rate during fermentation using the onggi's gas permeability as a free parameter. Our model provides a good fit for the data, and we conclude that porous walls help the onggi to 'exhale' carbon dioxide, lowering internal levels to those favoured by lactic acid bacteria. The positive pressure inside the onggi and the constant outflow through its walls act as a safety valve for bacteria growth by blocking the entry of external contaminants without mechanical components. We hope this study draws attention to the work of traditional artisans and inspires energy-efficient methods for fermenting and storing food products.

Canine-inspired Unidirectional Flows for Improving Memory Effects in Machine Olfaction.

A dog's nose differs from a human's in that air does not change direction but flows in a unidirectional path from inlet to outlet. Previous simulations showed that unidirectional flow through a dog's complex nasal passageways creates stagnant zones of trapped air. We hypothesize that these zones give the dog a "physical memory," which it may use to compare recent odors to past ones. In this study, we conducted experiments with our previously built Gaseous Recognition Oscillatory Machine Integrating Technology (GROMIT) and performed corresponding simulations in two dimensions. We compared three settings: a control setting that mimics the bidirectional flow of the human nose; a short-circuit setting where odors exit before reaching the sensors; and a unidirectional configuration using a dedicated inlet and outlet that mimics the dog's nose. After exposure to odors, the sensors in the unidirectional setting showed the slowest return to their baseline level, indicative of memory effects. Simulations showed that both short-circuit and unidirectional flows created trapped recirculation zones, which slowed the release of odors from the chamber. In the future, memory effects such as the ones found here may improve the sensitivity and utility of electronic noses.

ForageFeeder: A low-cost open source feeder for randomly distributing food.

Automated feeders have long fed mice, livestock, and poultry, but are incapable of feeding zoo animals such as gorillas. In captivity, gorillas eat cut vegetables and fruits in pieces too large to be dispensed by automated feeders. Consequently, captive gorillas are fed manually at set times and locations, keeping them from the exercise and enrichment that accompanies natural foraging. We designed and built ForageFeeder, an automated gorilla feeder that spreads food at random intervals throughout the day. ForageFeeder is an open source and easy to manufacture and modify device, making the feeder more accessible for zoos. The design presented here reduces manual labor for zoo staff and may be a useful tool for studies of animal ethology.