Seasonal Variations in the Toughness of Leaves: A Case Study Using Griselinia littoralis.

Potential effects of climate change include greater extremes of temperature and increased severity of storms. Many plants have evolved to resist the challenges of winter (freezing, dehydration, and wind) in a process known as cold hardening. Sensing reducing temperatures, they make structural changes at the cellular level to increase their mechanical resistance and prevent damage. Previous work on this topic, though extensive, has been conducted under laboratory conditions rather than in the field, and while many workers have observed changes to cell wall thickness and composition, which imply increased mechanical strength, few have actually measured strength or any other parameter describing structural integrity. This paper describes experiments on a model system designed to measure the structural integrity of leaf laminae from plants growing naturally in the field over extended periods, allowing seasonal variations to be captured. Standard engineering properties-tensile strength and fracture toughness-were measured for leaves of Griselinia littoralis on 19 separate occasions over a 12-month period. Toughness (rather than strength) was found to be the controlling mechanical property. Toughness values were found to change significantly during the year, by more than a factor of 2. Toughness correlated strongly with average daily soil temperature, but with a lag of about 1-2 weeks, suggesting that this is the time needed for structural adjustments to take place. Highest toughness values occurred in winter, confirming cold hardening. Increasing temperature in the spring was associated with decreasing toughness, but in the summer, when highest temperatures occurred, toughness increased again. This apparent "hot hardening" may be a response to dehydration. Results imply that a given leaf is able to both increase and decrease its toughness in response to temperature changes, demonstrating excellent plasticity of response. This case study of a single species establishes a method of reliably measuring changes in a plant's structural integrity due to cold hardening and other seasonal variations, which may be used to investigate the effects of climate change and other variables.