Measurement of leaf biomechanical properties in studies of herbivory: Opportunities, problems and procedures

Leaf biomechanical properties have the potential to act as antiherbivore defences. However, compared with studies on chemical defences, there are few studies that have demonstrated that the physical or biomechanical structure of plants can prevent or influence herbivory. This difference in focus by ecologists may relate to the dominant paradigm of plant chemical defences in ecological research and the perceived difficulties that ecologists have with the engineering principles embedded in biomechanics. The advantage of using materials engineering concepts is that each property is precisely defined and quantifiable, although the latter may be difficult in leaves because of their composite and anisotropic nature. Most herbivory studies have used simple penetrometers to measure leaf properties, often termed 'toughness'. As defined in materials engineering, the measured properties are 'force to fracture' and 'strength', not toughness. Measurement of strength, the resistance to crack initiation, is relevant to understanding herbivory. Measurement of 'toughness' as defined by materials engineering is also relevant. Toughness is the resistance to crack propagation and is a measure of the energy required to fracture the leaf. This requires more sophisticated equipment than simple penetrometers because it requires a simultaneous measure of the punch displacement. In addition, purists would argue that a punch cannot be used to measure true toughness because the crack is not controlled and plastic deformation is also involved. However, it may be the only method that allows detection of fine-scale pattern in mechanical properties across a leaf surface at a scale that is relevant to herbivory. There is very little work on the scale at which these properties vary, particularly with regard to different sized herbivores. In addition, few studies have investigated a broad range of relevant biomechanical properties in relation to herbivory. Therefore, it is not possible yet to be definitive about the relative merits of the various types of tests. A single test might show a pattern in relation to herbivore damage at a gross level. However, to really understand the functional and ecological significance of leaf texture in relation to herbivory, a more reductionist approach is needed. Only then can we move on to the larger scales of pattern that many ecologists are seeking.