Changes in spinescence across leaf ontogeny support the optimal defence hypothesis in blackberries (Rubus adenotrichos)

Abstract

Hypotheses based on allocation theory and herbivore selection offer opposite predictions about how defence levels against herbivores change as the plant tissue grows. The growth differentiation balance hypothesis (GDBH) assumes that defences will be resource-limited in immature tissues and predict that defence levels increase as the plant tissue grows. Conversely, the optimal defence hypothesis (ODH) proposes that plants would have the highest level of defences in the parts that have the highest value in terms of fitness and/or are more frequently attacked by herbivores, such as young tissues. We examine whether spinescence in the shrub Rubus adenotrichos (blackberry) change as the leaf grows, and if this change is consistent with the GDBH or the ODH. We compare the petiole area occupied by prickles, the prickles density and the individual prickle area in mature versus young petioles from Rubus adenotrichos. Our results show that, in R. adenotrichos, young tissues are more protected than mature tissues. Prickles density and the petiole area occupied by prickles were up to 25% higher in young petioles than in mature ones. These results support the ODH, reinforcing the idea that extrinsic factors such as herbivores pressure might drive the change of structural defences level across leaf ontogeny.