Sunflecks in the upper canopy: dynamics of light-use efficiency in sun and shade leaves of Fagus sylvatica.

Sunflecks are transient patches of direct radiation that provide a substantial proportion of the daily irradiance to leaves in the lower canopy. In this position, faster photosynthetic induction would allow for higher sunfleck-use efficiency, as is commonly reported in the literature. Yet, when sunflecks are too few and far between, it may be more beneficial for shade leaves to prioritize efficient photosynthesis under shade. We investigated the temporal dynamics of photosynthetic induction, recovery under shade, and stomatal movement during a sunfleck, in sun and shade leaves of Fagus sylvatica from three provenances of contrasting origin. We found that shade leaves complete full induction in a shorter time than sun leaves, but that sun leaves respond faster than shade leaves due to their much larger amplitude of induction. The core-range provenance achieved faster stomatal opening in shade leaves, which may allow for better sunfleck-use efficiency in denser canopies and lower canopy positions. Our findings represent a paradigm shift for future research into light fluctuations in canopies, drawing attention to the ubiquitous importance of sunflecks for photosynthesis, not only in lower-canopy leaves where shade is prevalent, but particularly in the upper canopy where longer sunflecks are more common due to canopy openness.

The Leaf Economics Spectrum Constrains Phenotypic Plasticity Across a Light Gradient.

The leaf economics spectrum (LES) characterizes multivariate correlations that confine the global diversity of leaf functional traits onto a single axis of variation. Although LES is well established for traits of sun leaves, it is unclear how well LES characterizes the diversity of traits for shade leaves. Here, we evaluate LES using the sun and shade leaves of 75 woody species sampled at the extremes of a within-canopy light gradient in a subtropical forest. Shading significantly decreased the mean values of LMA and the rates of photosynthesis and dark respiration, but had no discernable effect on nitrogen and phosphorus content. Sun and shade leaves manifested the same relationships among N mass, P mass, A mass, and R mass (i.e., the slopes of log-log scaling relations of LES traits did not differ between sun and shade leaves). However, the difference between the normalization constants of shade and sun leaves was correlated with functional trait plasticity. Although the generality of this finding should be evaluated further using larger datasets comprising more phylogenetically diverse taxa and biomes, these findings support a unified LES across shade as well as sun leaves.

Leaf functional trait variation in a humid temperate forest, and relationships with juvenile tree light requirements.

The species-rich arborescent assemblages of humid tropical forests encompass much of the known range of the leaf economics spectrum, often including >20-fold variation in leaf lifespan. This suite of traits underpins a life-history continuum from fast-growing pioneers to slow-growing shade-tolerant species. Less is known about the range of leaf traits in humid temperate forests, and there are conflicting reports about relationships of these traits with the light requirements of temperate evergreen angiosperms. Here I quantify the range of leaf functional traits in a New Zealand temperate evergreen forest, and relationships of these traits with light requirements of juvenile trees and shrubs. Foliage turnover of saplings of 19 evergreen angiosperms growing beneath gaps (12-29% canopy openness) and in understories (1.2-2.9%) was measured over 12 months. Dry mass per area (LMA), dry matter content, thickness, density and nitrogen content (N) of leaves were also measured. Species minimum light requirements were indexed as the 10th percentile of the distribution of saplings in relation to canopy openness. Interspecific variation of leaf lifespan was ∼6-fold in gaps (0.6 to 3.8 yrs), and ∼11-fold in the understorey (0.7 to 7.7 yrs). Six small tree and shrub species are effectively leaf-exchangers, with leaf lifespans of c.1 year in gaps-albeit usually longer in the shade. Interspecific variation in other leaf traits was 2.5 to 4-fold. Lifespans and LMA of both sun and shade leaves were negatively correlated with species light requirements i.e., positively correlated with shade tolerance. However, light environment (gap vs shade) explained about the same amount of variation in LMA as species' identity did. Species light requirements were not significantly correlated with leaf N, dry matter content, density or thickness-except for a marginally significant correlation with dry matter content of shade leaves. Species light requirements were thus less consistently related to leaf structural traits than appears to be the case in humid tropical forests. Whereas the wide interspecific variation in leaf economic traits of tropical rainforest species outweighs plastic response to light availability, temperate evergreen woody angiosperms appear to occupy a narrower range of the leaf economic spectrum. Standardization of the light environments in which LMA is measured is vital in comparative studies of humid temperate forest evergreens, because of countergradient responses of this trait to light, and because of the relative magnitudes of plastic and interspecific variation in LMA in these forests.

Low leaf-level response to light and nutrients in Mediterranean evergreen oaks: a conservative resource-use strategy?

We have explored leaf-level plastic response to light and nutrients of Quercus ilex and Q. coccifera, two closely related Mediterranean evergreen sclerophylls, in a factorial experiment with seedlings. Leaf phenotypic plasticity, assessed by a relative index (PI = (maximum value - minimum)/maximum) in combination with the significance of the difference among means, was studied in 37 morphological and physiological variables. Light had significant effects on most variables relating to photosynthetic pigments, chlorophyll fluorescence and gas exchange, whereas nutrient treatment had a significant effect in only 10% of the variables. Chlorophyll content was higher in the shade whereas carotenoid content and nonphotochemical quenching increased with light. Nutrient limitations increased the xanthophyll-cycle pool but only at high light intensities, and the same interaction between light and nutrients was observed for lutein. Predawn photochemical efficiency of PSII was not affected by either light or nutrients, although midday photochemical efficiency of PSII was lower at high light intensities. Photosynthetic light compensation point and dark respiration on an area basis decreased with light, but photosynthetic capacity on a dry mass basis and photochemical quenching were higher in low light, which translated into a higher nitrogen use efficiency in the shade. We expected Q. ilex, the species of the widest ecological distribution, to be more plastic than Q. coccifera, but differences were minor: Q. ilex exhibited a significant response to light in 13% more of the variables than Q. coccifera, but mean PI was very similar in the two species. Both species tolerated full sunlight and moderate shade, but exhibited a reduced capacity to enhance photosynthetic utilization of high irradiance. When compared with evergreen shrubs from the tropical rainforest, leaf responsiveness of the two evergreen oaks was low. We suggest that the low leaf-level responsiveness found here is part of a conservative resource use strategy, which seems to be adaptive for evergreen woody plants in Mediterranean-type ecosystems.

Is acclimation required for success in high light environments? A case study using Mycelis muralis (L.) Dumort (Asteraceae).

In the field significant differences in maximum photosynthetic O2 -exchange rate (Pm ) were found between leaves of Mycelis muralis (L.) Dumort (Asteraceae) collected from woodland and exposed habitats, with the highest values in the exposed sites- However, there were no differences in the Pm of leaves collected from plants growing in grikes (fissures in the limestone pavement), of exposed limestone pavement, despite a greater than four-fold difference in the integrated daily irradiance. Leaves of plants from the open pavement had lower photon yields (ø1 ) and higher dark respiration rates and light compensation points, in comparison to shaded plants. Under controlled environmental conditions the highest Pm of leaves from plants subjected to variations in irradiance were found at the intermediate (8-6 mol photon m-2 d-1 growth light level used. At the highest growth irradiance 17.3 mol photon m-2 d-1 used in the laboratory both Pm and øl were reduced, although the latest plant biomass was found at this irradiance. No changes were found in the chlorophyll a:b ratio over the same range of irradiances. Examination of plant populations of M. muralis, collected from open or shaded habitats and exposed to growth irradiances that covered the range over which increases in photosynthesis were, observed in the laboratory (0.86-8.6 mol photon m-2 d-1 ), resulted in changes in leaf structure and pigment composition. The chlorophyll a:b ratio was low and largely independent of irradiance or the origin of the plant population. Differences in total chlorophyll content were small with the lowest values m the Durrow woodland populations at both irradiances. No variations were found in a number of chloroplast thylakoid structural features. In particular, the ratio of oppressed to non-appressed membranes was unchanged by growth at the two irradiances, consistent with an invariant chlorophyll a:b ratio. Based on peaks in the difference spectra the woodland populations had mi enhanced in vivo absorption at λlD= 650 and 706 nm when grown at low irradiance. These peaks were absent from the population collected from the open limestone pavement. The significance of the enhanced absorption at low irradiance and the possibility that these peaks represent long-wavelength forms of chlorophyll a (λlD = 706) and b (λlD = 650) is discussed. A particular feature of plants grown at high irradiance was an enhanced anthocyanin content in comparison to those grown at low irradiance. This was associated with an increase in absorptance. particularly in the green region (λlD = 550 nm) of the visible spectrum. Overall these results suggest that complete acclimation of photosynthesis and an ability to modulate light-harvesting is not a prerequisite, for success in a high light environment.

Plasticity, instability and canalization: is the phenotypic variation in seedlings of sclerophyll oaks consistent with the environmental unpredictability of Mediterranean ecosystems?

• Evergreen oaks from the Mediterranean basin exhibit a conservative resource-use strategy based on a reduced expression of phenotypic variation (i.e. canalization). We hypothesized that genetic variation across closely related species is more canalized than the response to environmental variation. • Seedlings of Quercus ilex and Q. coccifera, two important oak species from the Mediterranean basin that belong to the same subgenus and section, were grown in contrasted light and nutrient environments following a factorial design. Phenotypic variation was explored in a total of 75 variables including photosynthetic capacity, nutrient allocation, allometric relationships and crown architecture. • Path analysis showed that phenotypic variation was not significantly affected by differences between species but by those between and within environments, which are argued to be primarily linked to phenotypic plasticity and developmental instability, respectively. This finding is interpreted as evidence of genetic canalization across species. • The similar importance of plasticity and instability as sources of phenotypic variation and the high degree of genetic canalization are consistent with the expected role of the environmental unpredictability of Mediterranean ecosystems in shaping the developmental patterns of these two species.