The competitive equilibrium between deciduous and evergreen plant species to a large extent depends on the intensity of the reduction in carbon gain undergone by evergreen leaves, associated with the leaf traits that confer resistance to stressful conditions during the unfavourable part of the year. This study explores the effects of winter harshness on the resistance traits of evergreen leaves. Leaf mass per unit area (LMA), leaf thickness and the concentrations of fibre, nitrogen (N), phosphorus (P), soluble protein, chlorophyll and ribulose 1,5-bisphosphate carboxylase/oxygenase (Rubisco) were determined in three evergreen and two deciduous species along a winter temperature gradient. In the evergreen species, LMA, thickness, and P and structural carbohydrate concentrations increased with the decrease in winter temperatures. Nitrogen and lignin concentrations did not show definite patterns in this regard. Chlorophyll, soluble proteins and Rubisco decreased with the increase in winter harshness. Our results suggest that an increase in LMA and in the concentration of structural carbohydrates would be a requirement for the leaves to cope with low winter temperatures. The evergreen habit would be associated with higher costs at cooler sites, because the cold resistance traits imply additional maintenance costs and reduced N allocation to the photosynthetic machinery, associated with structural reinforcement at colder sites.
Studies on plasticity at the level of a single individual plant provide indispensable information to predict leaf responses to climate change, because they allow better identification of the environmental factors that determine differences in leaf traits in the absence of genetic differences. Most of these studies have focused on the responses of leaf traits to variations in the light environment along vertical gradients, thus paying less attention to possible differences in the intensity of water stress among canopy orientations. In this paper, we analyzed the differences in leaf traits traditionally associated with changes in the intensity of water stress between east and west crown orientations in three Quercus species. The leaves facing west experienced similar solar radiation levels but higher maximum temperatures and lower daily minimum water potentials than those of the east orientation. In response to these differences, the leaves of the west orientation showed smaller size and less chlorophyll concentration, higher percentage of palisade tissue and higher density of stomata and trichomes. These responses would confirm the role of such traits in the tolerance to water stress and control of water losses by transpiration. For all traits, the species with the longest leaf life span exhibited the greatest plasticity between orientations. By contrast, no differences between canopy positions were observed for leaf thickness, leaf mass per unit area and venation patterns.
Investment in anti‐herbivore defence in tree species has been one of the priority research topics in plant terrestrial ecology during the last decades. However, despite considerable experimental effort, interspecific differences in the ontogenetic trends in the investment in defence are still a matter of debate, as to date experimental evidence is contradictory. In the present work, insect herbivory levels were measured in seedlings and mature trees of four co‐occurring Mediterranean Quercus species with differing leaf life spans, as well as several leaf characteristics that can determine herbivore preference. The measured leaf traits included nitrogen (N), fibre (cellulose, hemicellulose and lignin), total phenolic contents, leaf mass per unit area (LMA) and leaf thickness. The leaves of seedlings had a lower LMA and leaf thickness and lower concentrations of N and cellulose, but higher concentrations of lignin and phenols than those of mature trees. However, the loss of leaf area tended to be more severe for seedlings than for mature trees, although the differences were only significant for deciduous species. This constitutes a confirmation of the strong effects of physical traits on herbivore preferences. The greater resource limitations for defensive mechanisms in seedlings with respect to mature trees would explain that at intraspecific level we do observe a compromise between chemical and physical defences. As a result, seedlings rely on chemical rather than on physical defences.
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