Novel developments in field mechanics.

Our aim is general: we want to illustrate how much can be gleaned from mechanical measurement in the field. We ask how mechanics may constrain foraging and feeding on both plants and animals, and how various aspects of mechanical behavior could affect the feeding choices that primates make. Here, we present novel methods for the measurement of the material properties and also the employment of tried and tested methods in novel settings. This review demonstrates how mechanical investigation methods can quantify the environmental factors affecting primate locomotion to and from food, which makes up a large part of a primate's daily energy budget. We indicate that, despite the accumulation of much data on the material properties of primate foods, the introduction of new methods is allowing researchers to pursue new avenues of research and change paradigms in primate feeding ecology. Field methods are presented that could aid in the understanding of the extra-oral processing of foodstuffs by primates and enrich further studies into cognition and culture surrounding these types of behavior. We conclude that the use of in-field measurements and a greater understanding of the physics of primate environments are vital and exciting themes integral to the continued understanding of primate evolution and biology.

Nest-building orangutans demonstrate engineering know-how to produce safe, comfortable beds.

Nest-building orangutans must daily build safe and comfortable nest structures in the forest canopy and do this quickly and effectively using the branches that surround them. This study aimed to investigate the mechanical design and architecture of orangutan nests and determine the degree of technical sophistication used in their construction. We measured the whole nest compliance and the thickness of the branches used and recorded the ways in which the branches were fractured. Branch samples were also collected from the nests and subjected to three-point bending tests to determine their mechanical properties. We demonstrated that the center of the nest is more compliant than the edges; this may add extra comfort and safety to the structure. During construction orangutans use the fact that branches only break half-way across in "greenstick" fracture to weave the main nest structure. They choose thicker branches with greater rigidity and strength to build the main structure in this way. They then detach thinner branches by following greenstick fracture with a twisting action to make the lining. These results suggest that orangutans exhibit a degree of technical knowledge and choice in the construction of nests.

Factors affecting the compliance and sway properties of tree branches used by the Sumatran orangutan (Pongo abelii).

The tropical arboreal environment is a mechanically complex and varied habitat. Arboreal inhabitants must adapt to changes in the compliance and stability of supports when moving around trees. Because the orangutan is the largest habitual arboreal inhabitant, it is unusually susceptible to branch compliance and stability and therefore represents a unique animal model to help investigate how animals cope with the mechanical heterogeneity of the tropical canopy. The aim of this study was to investigate how changes in compliance and time of oscillation of branches are related to easily observable traits of arboreal supports. This should help predict how supports react mechanically to the weight and mass of a moving orangutan, and suggest how orangutans themselves predict branch properties. We measured the compliance and time of oscillation of branches from 11 tree species frequented by orangutans in the rainforest of Sumatra. Branches were pulled at several points along their length using a force balance at the end of a stiff rope, and the local diameter of the branch and the distance to its base and tip were measured. Compliance was negatively associated with both local diameter and length to the tip of the branch, and positively, if weakly, associated with length from the trunk. However, branch diameter not only predicted compliance best, but would also be easiest for an orangutan to observe. In contrast, oscillation times of branches were largely unaffected by local diameter, and only significantly increased at diameters below 2 cm. The results of this study validate previous field research, which related locomotory modes to local branch diameter, while suggesting how arboreal animals themselves sense their mechanical environment.

Fingerprints are unlikely to increase the friction of primate fingerpads.

It is generally assumed that fingerprints improve the grip of primates, but the efficiency of their ridging will depend on the type of frictional behaviour the skin exhibits. Ridges would be effective at increasing friction for hard materials, but in a rubbery material they would reduce friction because they would reduce contact area. In this study we investigated the frictional performance of human fingertips on dry acrylic glass using a modified universal mechanical testing machine, measuring friction at a range of normal loads while also measuring the contact area. Tests were carried out on different fingers, fingers at different angles and against different widths of acrylic sheet to separate the effects of normal force and contact area. The results showed that fingertips behaved more like rubbers than hard solids; their coefficients of friction fell at higher normal forces and friction was higher when fingers were held flatter against wider sheets and hence when contact area was greater. The shear stress was greater at higher pressures, suggesting the presence of a biofilm between the skin and the surface. Fingerprints reduced contact area by a factor of one-third compared with flat skin, however, which would have reduced the friction; this casts severe doubt on their supposed frictional function.

The effect of humidity on the fracture properties of human fingernails.

Fingernails are a characteristic anatomical feature of primates and their function is dictated by the environment in which they are utilised. The present study examined the mechanical properties of human fingernails as a function of relative humidity (RH) and the subsequent moisture content of the nail material. Nail clippings were stored at a range of RH values and then weighed in order to determine their moisture content. There was a non-linear relationship between the moisture content of nails and the RH of their local environment. The in vivo moisture content of nails, measured from 55% to 80% RH, was between 14% and 30%, similar to other keratinous materials such as claws, hooves and feathers. Cutting tests on the nail samples showed that the work of fracture was between 11 and 22 kJ m(-2), rising to a peak at 55% RH and falling at higher and lower humidities. At all RH values there was anisotropy within the nail between the proximal and lateral directions, the work of fracture being greater proximally. This anisotropy was greatest at 55% RH, at which point the proximal work of fracture was double the lateral value. These results suggest that the mechanical behaviour of human fingernails is optimised at in vivo conditions; they resist tearing most strongly under these conditions and particularly resist tearing into the nail bed. At more extreme humidity levels the fracture properties of the nail deteriorate; they are brittle when fully dry and fracture and split when wet.

Interrelations between hydraulic and mechanical stress adaptations in woody plants.

The fields of plant water relations and plant biomechanics have traditionally been studied separately even though often the same tissues are responsible for water transport and mechanical support. There is now increasing evidence that hydraulic and mechanical adaptations may influence one another. We studied the changes in the hydraulic and mechanical properties of the wood along lateral roots of two species of buttressed trees. In these roots, the mechanical contstraints quantified by strain measurements are known to decrease distally. Further, we investigated the effect of mechanical loading on the vessel anatomy in these and four other species of tropical trees. We found that as the strain decreased, the wood became progressively less stiff and strong but the conductivity increased exponentially. This was reflected in that adaptations towards re-enforcing mechanically loaded areas resulted in xylem with fewer and smaller vessels. In addition a controlled growth experiment on three tree species showed that drought adaptation may results in plants with stronger and stiffer tissue. Our results indicate that hydraulic and mechanical stress adaptations may be interrelated, and so support recent studied suggesting that physiological responses are complex balances rather than pure optimisations.

Herbivore specific induction of silica-based plant defences.

Induced plant responses to herbivory have major impacts on herbivore feeding behaviour, performance and population dynamics. These effects are well established for chemical defences, but induction of physical defences remains far less studied. However, for many plants, it is physical defences that play the major role in regulating the levels of herbivore damage sustained. We provide evidence that, in grasses, induction of physical defences is both specific to herbivore feeding, as opposed to mechanical damage, and may be dependant on the amount of damage imposed. Furthermore, we show that the magnitude of the induction response is sufficient to deter further damage and affect herbivore performance. We compared silica induction in two grass species in response to vertebrate and invertebrate damage, and to mechanical defoliation. Induction was assessed at two levels of damage over 16 months. Foliar silica content did not increase in response to mechanical defoliation, but damage by either voles or locusts resulted in increases in silica content of over 400%. This increase deterred feeding by both voles and locusts. Silica induction in grasses due to repeated damage events over a prolonged period suggests a possible role for silica defence in the cyclical population fluctuations observed in many grass-feeding herbivores.

Functional significance of the uncinate processes in birds.

The functional significance of the uncinate processes to the ventilatory mechanics of birds was examined by combining analytical modeling with morphological techniques. A geometric model was derived to determine the function of the uncinate processes and relate their action to morphological differences associated with locomotor specializations. The model demonstrates that uncinates act as levers, which improve the mechanical advantage for the forward rotation of the dorsal ribs and therefore lowering of the sternum during respiration. The length of these processes is functionally important; longer uncinate processes increasing the mechanical advantage of the Mm. appendicocostales muscle during inspiration. Morphological studies of four bird species showed that the uncinate process increased the mechanical advantage by factors of 2-4. Using canonical variate analysis and analysis of variance we then examined the variation in skeletal parameters in birds with different primary modes of locomotion (non-specialists, walking and diving). Birds clustered together in distinct groups, indicating that uncinate length is more similar in birds that have the same functional constraint, i.e. specialization to a locomotor mode. Uncinate processes are short in walking birds, long in diving species and of intermediate length in non-specialist birds. These results demonstrate that differences in the breathing mechanics of birds may be linked to the morphological adaptations of the ribs and rib cage associated with different modes of locomotion.

Silica in grasses as a defence against insect herbivores: contrasting effects on folivores and a phloem feeder.

1. Silica, deposited as opaline phytoliths in the leaves of grasses, constitutes 2-5% of dry leaf mass, yet its function remains unclear. It has been proposed that silica may act as an antiherbivore defence by increasing the abrasiveness and reducing the digestibility of grass leaves, although there is little direct experimental evidence to support this. 2. We investigated the effects of manipulated silica levels on the abrasiveness of the leaves of five grass species. We also examined the effects of silica levels on the feeding preferences, growth performance and digestion efficiency of two folivorous insects and one phloem-feeding insect. 3. Silica addition resulted in increases to leaf abrasiveness in four of the five grass species studied. Silica addition also deterred feeding by both folivores and reduced their growth rates and digestion efficiency. 4. These effects resulted in lower pupal mass of the lepidopteron larvae Spodoptera exempta and compensatory feeding by the orthopteran, Schistocerca gregaria. In contrast, silica had no effects on the feeding preference or the population growth of the phloem feeder, Sitobion avenae. 5. Our results demonstrate that silica is an effective defence against folivorous insects, both as a feeding deterrent, possibly mediated by increased abrasiveness, and as a digestibility reducer. The effects of silica on pupal mass and development time may impact on herbivore fitness and exposure to natural enemies. 6. These results are the first demonstration of a direct effect of silica on the abrasiveness of grasses and the adverse impact of silica on herbivore preference and performance.