The biomechanics of browsing and grazing

Terrestrial plant leaves are complex structures of composite materials. Resistance to fracture is achieved by a number of mechanisms, which operate at the molecular, cell, tissue, and structural levels. Leaves of dicots have different venation patterns and cell wall volume fractions from those of grasses, and consequently, they potentially resist fracture in different ways. Animals mechanically process plants in order to rupture the cell wall in preparation for enzymic hydrolysis, for which the imperative is to maximize new surface area and/or to expedite access to cell contents, ideally by promoting elastic fracture. The two different plant types are fed on by two different groups of organisms of very different sizes, digestive physiologies, mechanical processing abilities and properties, and nutritional requirements. Small insects can feed in or on parts of leaves, while larger mammals generally have to feed on the whole leaf. The scale of feeding also differs for the two groups of herbivores, but how this interacts with the scale of the mechanical properties of the leaf is not well understood. Plant leaves are attacked at all scales and probably can only produce generalized responses to specialized herbivores. In addition, the opportunities that these different scales of interactions open for the different herbivores remain unexplored.