What does 'toughness' look like? An examination of the breakdown of young and mature leaves under cyclical loading.

The material property of leaf toughness is considered the crucial mechanical challenge facing folivorous primates. Mature leaves have higher recorded toughness values than young leaves on average, leading to many assumptions about the patterning of food breakdown that follow a tough/not-tough dichotomy. We tested three hypotheses about how leaves break down under repetitive loading cycles, predicting that mature leaves (i) experience more force during simulated occlusal loads, (ii) more effectively resist fragmentation into small pieces, and (iii) show a more gradual decline in resistance over consecutive cycles than young leaves. Under displacement control using a mechanical testing system, we subjected young and mature leaves to 20 cycles of axial loading using interlocking steel wedges, then collected and quantified the size of the leaf fragments. While we found that mature leaves experienced more overall force than young leaves (p < 0.001), they also shattered into smaller pieces (p = 0.004) and showed a steeper decline in their resistance to the cycles over the course of a test (p < 0.01). These results suggest that putatively 'tougher' foods (i.e. mature versus young leaves) do not necessarily resist fragmentation as commonly assumed. The current tough/not-tough paradigm of primate foods may not accurately reflect how leaves break down during masticatory behaviour.