Br-rich tips of calcified crab claws are less hard but more fracture resistant: a comparison of mineralized and heavy-element biological materials.

We find that the spoon-like tips of the chelipeds (large claws) of the crab Pachygrapsus crassipes differ from the rest of the claw in that they are not calcified, but instead contain about 1% bromine--thus they represent a new example of a class of structural biological materials that contain heavy elements such as Zn, Mn, Fe, Cu, and Br bound in an organic matrix. X-ray absorption spectroscopy data suggest that the bromine is bound to phenyl rings, possibly in tyrosine. We measure a broad array of mechanical properties of a heavy-element biological material for the first time (abrasion resistance, coefficient of kinetic friction, energy of fracture, hardness, modulus of elasticity and dynamic mechanical properties), and we make a direct comparison with a mineralized tissue. Our results suggest that the greatest advantage of bromine-rich cuticle over calcified cuticle is resistance to fracture (the energy of fracture is about an order of magnitude greater than for calcified cuticle). The greatest advantage relative to unenriched cuticle, represented by ant mandible cuticle, is a factor of about 1.5 greater hardness and modulus of elasticity.The spoon-like tips gain additional fracture resistance from the orientation of the constituent laminae and from the viscoelasticity of the material. We suggest that fracture resistance is of greater importance in smaller organisms, and we speculate that one function of heavy elements in structural biological materials is to reduce molecular resonant frequencies and thereby increase absorption of energy from impacts.

Leaf processing behaviour in Atta leafcutter ants: 90% of leaf cutting takes place inside the nest, and ants select pieces that require less cutting.

Leafcutter ants cut trimmings from plants, carry them to their underground nests and cut them into smaller pieces before inoculating them with a fungus that serves as a primary food source for the colony. Cutting is energetically costly, so the amount of cutting is important in understanding foraging energetics. Estimates of the cutting density, metres of cutting per square metre of leaf, were made from samples of transported leaf cuttings and of fungal substrate from field colonies of Atta cephalotes and Atta colombica. To investigate cutting inside the nest, we made leaf-processing observations of our laboratory colony, A. cephalotes. We did not observe the commonly reported reduction of the leaf fragments into a pulp, which would greatly increase the energy cost of processing. Video clips of processing behaviours, including behaviours that have not previously been described, are linked. An estimated 2.9 (±0.3) km of cutting with mandibles was required to reduce a square metre of leaf to fungal substrate. Only about 12% (±1%) of this cutting took place outside of the nest. The cutting density and energy cost is lower for leaf material with higher ratios of perimeter to area, so we tested for, and found that the laboratory ants had a preference for leaves that were pre-cut into smaller pieces. Estimates suggest that the energy required to transport and cut up the leaf material is comparable to the metabolic energy available from the fungus grown on the leaves, and so conservation of energy is likely to be a particularly strong selective pressure for leafcutter ants.

Is fracture a bigger problem for smaller animals? Force and fracture scaling for a simple model of cutting, puncture and crushing.

Many of the materials that are challenging for large animals to cut or puncture are also cut and punctured by much smaller organisms that are limited to much smaller forces. Small organisms can overcome their force limitations by using sharper tools, but one drawback may be an increased susceptibility to fracture. We use simple contact mechanics models to estimate how much smaller the diameter of the tips or edges of tools such as teeth, claws and cutting blades must be in smaller organisms in order for them to puncture or cut the same materials as larger organisms. In order to produce the same maximum stress when maximum force scales as the square of body length, the diameter of the tool region that is in contact with the target material must scale isometrically for punch-like tools (e.g. scorpion stings) on thick targets, and for crushing tools (e.g. molars). For punch-like tools on thin targets, and for cutting blades on thick targets, the tip or edge diameters must be even smaller than expected from isometry in smaller animals. The diameters of a small sample of unworn punch-like tools from a large range of animal sizes are consistent with the model, scaling isometrically or more steeply (positively allometric). In addition, we find that the force required to puncture a thin target using real biological tools scales linearly with tip diameter, as predicted by the model. We argue that, for smaller tools, the minimum energy to fracture the tool will be a greater fraction of the minimum energy required to puncture the target, making fracture more likely. Finally, energy stored in tool bending, relative to the energy to fracture the tool, increases rapidly with the aspect ratio (length/width), and we expect that smaller organisms often have to employ higher aspect ratio tools in order to puncture or cut to the required depth with available force. The extra stored energy in higher aspect ratio tools is likely to increase the probability of fracture. We discuss some of the implications of the suggested scaling rules and possible adaptations to compensate for fracture sensitivity in smaller organisms.

Tooth hardness increases with zinc-content in mandibles of young adult leaf-cutter ants.

A wide variety of arthropods and members of other phyla have elevated concentrations of Zn, Mn, other heavy metals and halogens in their jaws, leg claws, and other "tools" for interacting with the environment. While measured Zn concentrations reach 25% of dry mass in scorpion stings, concentrations are often lower than this and the enriched structures are not heavily biomineralized like vertebrate teeth and the radula of mollusks. For this reason, the degree to which the inorganic components of these structures modify their mechanical properties is in question. Here we address this problem by measuring hardness during the development of Zn accumulations in ant mandibles. We found that Zn is incorporated into the mandibular teeth of leaf-cutter ants during early adult life, reaching concentrations of about 16% of dry mass. We show that the hardness of the mandibular teeth increases nearly three-fold as the adults age and that hardness correlates with Zn content ( r=0.91). We suggest that young adults rarely cut leaves partly because their mandibles are not yet rich in Zn. Zinc enrichment (along with enrichment by other heavy metals and halogens) may play an unrecognized role in the behavioral ecology and evolution of a wide variety of invertebrates.