Why are evergreen leaves so contrary about shade?

Leaf mass per area (LMA) is one of the most widely measured of all plant functional traits. In deciduous forests, there is similarity between plastic and evolutionary responses of LMA to light gradients. In evergreens, however, LMA is lower in shaded than sunlit individuals of the same species, whereas shade-tolerant evergreens have higher LMA than light-demanders grown under the same conditions. We suggest that this pattern of 'counter-gradient variation' results from some combination of (i) close evolutionary coordination of LMA with leaf lifespan, (ii) selection for different leaf constitutions (relative investment in cell walls versus cell contents) in sun and shade environments and/or (iii) constraints on plasticity as a result of genetic correlations between phenotypes expressed in sun and shade.     

Links between leaf anatomy and leaf mass per area of herbaceous species across slope aspects in an eastern Tibetan subalpine meadow

Leaf anatomy varies with abiotic factors and is an important trait for understanding plant adaptive responses to environmental conditions. Leaf mass per area (LMA) is a key morphological trait and is related to leaf performance, such as light‐saturated photosynthetic rate per leaf mass, leaf mechanical strength, and leaf lifespan. LMA is the multiplicative product of leaf thickness (LT) and leaf density (LD), both of which vary with leaf anatomy. Nevertheless, how LMA, LT, and LD covary with leaf anatomy is largely unexplored along natural environmental gradients. Slope aspect is a topographic factor that underlies variations in solar irradiation, air temperature, humidity, and soil fertility. In the present study, we examined (1) how leaf anatomy varies with different slope aspects and (2) how leaf anatomy is related to LMA, LD, and LT. Leaf anatomy was measured for 30 herbaceous species across three slope aspects (south‐, west‐, and north‐facing slopes; hereafter, SFS, WFS, and NFS, respectively) in an eastern Tibetan subalpine meadow. For 18 of the 30 species, LMA data were available from previous studies. LD was calculated as LMA divided by LT. Among the slope aspects, the dominant species on the SFS exhibited the highest LTs with the thickest spongy mesophyll layers. The thicker spongy mesophyll layer was related to a lower LD via larger intercellular airspaces. In contrast, LD was the highest on NFS among the slope aspects. LMA was not significantly different among the slope aspects because higher LTs on SFS were effectively offset by lower LDs. These results suggest that the relationships between leaf anatomy and LMA were different among the slope aspects. Mechanisms underlying the variations in leaf anatomy may include different solar radiation, air temperatures, soil water, and nutrient availabilities among the slope aspects.     

Leaf Mechanical Properties in Sclerophyll Woodland and Shrubland on Contrasting Soils

Sclerophylly is a common feature of vegetation on infertile soils, and its adaptive significance has been linked to nutrient-use efficiency by protection of leaves to maximise carbon gain. However, there has been little investigation of how the leaf mechanical properties that contribute to the phenomenon of sclerophylly vary along nutrient gradients. In this paper, we investigate how leaf mechanical properties vary among plants on three contrasting soil types (grey sand, laterite soil, and soil overlying dolerite) in a Mediterranean climate in southwestern Australia. Most species were sclerophyllous, but there was 5-fold variation in leaf mass per unit area (LMA) and 17- to 473-fold variation in mechanical properties among species. Species growing on laterite and/or sand (low-nutrient soils) had higher punch strength, work (a measure of toughness) to punch, specific (per unit leaf thickness) work to punch, work to shear, specific work to shear, and flexural stiffness (EI _W) than those on dolerite soils (higher in nutrients). There were few differences in mean values of leaf mechanical properties between the two low-nutrient soils, possibly because the lower concentration of nutrients in the sand is balanced by the greater soil volume than the laterite soil (higher concentration of nutrients, but shallower). There were also few differences in leaf properties between plants of the same species growing on contrasting soil types. There was some variation among sclerophyllous species in their mechanical characteristics, but overall, EI _W provided the strongest contribution to sclerophylly, explaining up to 81% of the variation in LMA. There was no evidence of differences among soil types in the relationships of mechanical properties with LMA, and therefore, no evidence of variation in the mechanical constitution of sclerophylly among soil types.     

What makes a leaf tough? Patterns of correlated evolution between leaf toughness traits and demographic rates among 197 shade-tolerant woody species in a neotropical forest

Slow-growing juveniles of shade-tolerant plant species are predicted to have tough leaves because of the high cost of leaf replacement in shade relative to potential carbon gain. We assessed the degree of correlated evolution among eight traits associated with leaf toughness and the relationships of those traits with the growth and mortality rates of 197 tree and shrub species from the understory of the 50-ha forest dynamics plot on Barro Colorado Island, Panama. Path analysis with phylogenetically independent contrasts revealed that leaves attained material toughness (resistance to fracture per unit fracture area) through increases in tissue density, percent cellulose per unit dry mass, and vein fracture toughness. Lamina density and cellulose content evolved independently and thus represent different paths to material toughness. Structural toughness (resistance to fracture per unit fracture length) depended on material toughness and lamina thickness. Mortality rates of individuals 1-10 cm in stem diameter were negatively correlated with material toughness and lamina density but were independent of structural toughness and cell wall fiber contents. Leaf toughness traits were uncorrelated with relative growth rates. Results imply that material toughness enhances resistance to natural enemies, which increases survival and offsets the biomass allocation cost of producing tough leaves in the shaded understory.     

Contrasting foliar responses to drought in Quercus ilex and Phillyrea latifolia

Leaf morphology, longevity, and demography were examined in Quercus ilex and Phillyrea latifolia growing in a holm oak forest in Prades mountains (northeast Spain). Four plots (10 × 15 m) of this forest were submitted to an experimental drought during three years (soil moisture was reduced about 15 %). Leaf area, thickness and leaf mass per area ratio (LMA) were measured in sun and shade leaves of both species. Leaf longevity, the mean number of current-year shoots produced per previous-year shoot (Sn/Sn-1), the mean number of current-year leaves per previous-year shoot (Ln/Sn-1), and the percentage of previous-year shoots that developed new ones were measured once a year, just after leaf flushing. LMA and leaf thickness increased since leaf unfolding except in summer periods, when stomatal closure imposed low photosynthetic rates and leaves consumed their reserves. LMA, leaf area, and leaf thickness were higher in Q. ilex than in P. latifolia, but leaf density was higher in the latter species. Drought reduced the leaf thickness and the LMA of both species ca. 2.5 %. Drought also increased leaf shedding up to ca. 20 % in Phillyrea latifolia and decreased it up to ca. 20 % in Q. ilex. In the later species, Sn/Sn-1 decreased by 32 %, Ln/Sn-1 by 41 %, percentage of shoots developed new ones by 26 %, and leaf area by 17 %. Thus the decrease of leaf number and area was stronger in the less drought-resistant Q. ilex, which, under increasingly drier conditions, might lose its current competitive advantage in these Mediterranean holm oak forests.     

Leaf age and season influence the relationships between leaf nitrogen, leaf mass per area and photosynthesis in maple and oak trees

Abstract. Seasonal changes in photosynthesis, leaf nitrogen (N) contents and leaf mass per area (LMA) were observed over three growing seasons in open-grown sun-lit leaves of red maple ( Acer rubrum ), sugar maple ( A. sacchamm ) and northern pin oak ( Quereus ellipsoidalis ) trees in southern Wisconsin. Net photosynthesis and leaf N were highly linearly correlated on both mass and area bases within all species from late spring until leaf senescence in fall. Very early in the growing season leaves had high N concentrations, but low photosynthetic rates per unit leaf N, suggesting that leaves were not fully functionally developed at that time. Leaf N per unit area and LMA had nonparallel seasonal patterns, resulting in differing relationships between leaf N/area and LMA in the "early versus late growing season. As a result of differences in seasonal patterns between leaf N/area and LMA, net photosynthesis/area was higher for a given LMA in the spring than fall, and the overall relationships between these two parameters were poor.     

Influence of light level on the growth and morphology of saplings in a panamanian forest

Leaf spacing and aboveground growth were monitored in saplings of ten species in a range of light environments in a Panamanian lowland humid forest. One- to 2-m-tall individuals of the chosen species had intermediate to large leaves on stems with few or no branches. Saplings in high light environments grew faster in height and produced more biomass per unit leaf area than shaded saplings for all species. These growth responses involved morphological plasticity with greater extension per unit biomass increment increasing the height growth rate of gap-grown saplings and greater biomass allocation to leaves decreasing the whole plant light compensation point in shade. The relative performance of the species also varied across the light gradient and was related to differences in leaf lifespan and specific leaf mass. Light-demanding species grew as rapidly in shade as shade-tolerant species, but the shorter leaf lifespan of the former necessitates higher production rates to maintain a given leaf area, largely excluding light-demanders from shaded understory locations. Height growth rate was positively correlated with leaf spacing for each species, and differences between species in the height growth rate-internode length relationship were related to interspecific differences in specific leaf mass. Thus, sapling growth histories may be inferred from their morphologies.     

Leaf area and photosynthesis of newly emerged trifoliolate leaves are regulated by mature leaves in soybean

Leaf anatomy and the stomatal development of developing leaves of plants have been shown to be regulated by the same light environment as that of mature leaves, but no report has yet been written on whether such a long-distance signal from mature leaves regulates the total leaf area of newly emerged leaves. To explore this question, we created an investigation in which we collected data on the leaf area, leaf mass per area (LMA), leaf anatomy, cell size, cell number, gas exchange and soluble sugar content of leaves from three soybean varieties grown under full sunlight (NS), shaded mature leaves (MS) or whole plants grown in shade (WS). Our results show that MS or WS cause a marked decline both in leaf area and LMA in newly developing leaves. Leaf anatomy also showed characteristics of shade leaves with decreased leaf thickness, palisade tissue thickness, sponge tissue thickness, cell size and cell numbers. In addition, in the MS and WS treatments, newly developed leaves exhibited lower net photosynthetic rate (Pn), stomatal conductance (Gs) and transpiration rate (E), but higher carbon dioxide (CO ₂ ) concentration in the intercellular space (Ci) than plants grown in full sunlight. Moreover, soluble sugar content was significantly decreased in newly developed leaves in MS and WS treatments. These results clearly indicate that (1) leaf area, leaf anatomical structure, and photosynthetic function of newly developing leaves are regulated by a systemic irradiance signal from mature leaves; (2) decreased cell size and cell number are the major cause of smaller and thinner leaves in shade; and (3) sugars could possibly act as candidate signal substances to regulate leaf area systemically.     

How cellulose-based leaf toughness and lamina density contribute to long leaf lifespans of shade-tolerant species

Cell wall fibre and lamina density may interactively affect leaf toughness and leaf lifespan. Here, we tested this with seedlings of 24 neotropical tree species differing in shade tolerance and leaf lifespan under standardized field conditions (140-867 d in gaps; longer in shade). We quantified toughness with a cutting test, explicitly seeking a mechanistic linkage to fibre. Lamina density, but not fracture toughness, exhibited a plastic response to gaps vs shade, while neither trait was affected by leaf age. Toughness corrected for lamina density, a recently recognized indicator of material strength per unit mass, was linearly correlated with cellulose content per unit dry mass. Leaf lifespan was positively correlated with cellulose and toughness in shade-tolerant species but only weakly in gap-dependent species. Leaf lifespan was uncorrelated with lamina thickness, phenolics and tannin concentrations. In path analysis including all species, leaf lifespan was directly enhanced by density and toughness, and indirectly by cellulose via its effect on toughness. Different suites of leaf traits were correlated with early seedling survival in gaps vs shade. In conclusion, cellulose and lamina density jointly enhance leaf fracture toughness, and these carbon-based physical traits, rather than phenolic-based defence, explain species differences in herbivory, leaf lifespan and shade survival.     

Responses of leaf structure and photosynthetic properties to intra-canopy light gradients: a common garden test with four broadleaf deciduous angiosperm and seven evergreen conifer tree species

Spectra of leaf traits in northern temperate forest canopies reflect major differences in leaf longevity between evergreen conifers and deciduous broadleaf angiosperms, as well as plastic modifications caused by within-crown shading. We investigated (1) whether long-lived conifer leaves exhibit similar intra-canopy plasticity as short-lived broadleaves, and (2) whether global interspecific relationships between photosynthesis, nitrogen, and leaf structure identified for sun leaves adequately describe leaves differentiated in response to light gradients. We studied structural and photosynthetic properties of intra-tree sun and shade foliage in adult trees of seven conifer and four broadleaf angiosperm species in a common garden in Poland. Shade leaves exhibited lower leaf mass-per-area (LMA) than sun leaves; however, the relative difference was smaller in conifers than in broadleaves. In broadleaves, LMA was correlated with lamina thickness and tissue density, while in conifers, it was correlated with thickness but not density. In broadleaves, but not in conifers, reduction of lamina thickness was correlated with a thinner palisade layer. The more conservative adjustment of conifer leaves could result from a combination of phylogenetic constraints, contrasting leaf anatomies and shoot geometries, but also from functional requirements of long-lived foliage. Mass-based nitrogen concentration (N(mass)) was similar between sun and shade leaves, and was lower in conifers than in deciduous broadleaved species. Given this, the smaller LMA in shade corresponded with a lower area-based N concentration (N(area)). In evergreen conifers, LMA and N(area) were less powerful predictors of area-based photosynthetic rate (A (max(area))) in comparison with deciduous broadleaved angiosperms. Multiple regression for sun and shade leaves showed that, in each group, A (max(mass)) was related to N(mass) but not to LMA, whereas LMA became a significant codeterminant of A (max(mass)) in analysis combining both groups. Thus, a fundamental mass-based relationship between photosynthesis, nitrogen, and leaf structure reported previously also exists in a dataset combining within-crown and across-functional type variation.     

Is N‐resorption efficiency related to secondary compounds and leaf longevity in coexisting plant species of the arid Patagonian Monte,Argentina?

Leaf longevity and nutrient resorption efficiency are important strategies to conserve plant nutrients. Theory suggests a negative relationship between them and also proposes that high concentration of phenolics in long‐lived leaves may reduce nutrient resorption. In order to provide new evidence on these relationships, we explored whether N‐resorption efficiency is related to leaf longevity, secondary compounds and other leaf traits in coexisting plant species of different life forms in the arid Patagonian Monte, Argentina. We assessed N‐resorption efficiency, green leaf traits (leaf mass per area (LMA), leaf longevity and lignin, total soluble phenolics and N concentrations) and N concentration in senescent leaves of 12 species of different life forms (evergreen shrubs, deciduous shrubs and perennial grasses) with contrasting leaf traits. We found that leaf longevity was positively correlated to LMA and lignin, and negatively correlated to N concentration in green leaves. N concentrations both in green and senescent leaves were positively related. N‐resorption efficiency was not associated with the concentration of secondary compounds (total soluble phenolics and lignin) but it was negatively related to LMA and leaf longevity and positively related to N concentration in green leaves. Furthermore, leaf traits overlapped among life forms highlighting that life forms are not a good indicator of the functional properties (at least in relation to nutrient conservation) of species. In conclusion, our findings indicated that differences in N‐resorption efficiency among coexisting species were more related to N concentration in green leaves, leaf lifespan and LMA than to the presence of secondary compounds at least those assessed in our study (soluble phenolics and lignin). Accordingly, N‐resorption efficiency seems to be modulated, at least in part, by the productivity–persistence trade‐off.     

Herbivory pattern and induced responses to simulated herbivory inQuercus mongolica var.grosseserrata

Seasonal changes in leaf traits and the herbivory pattern ofQuercus mongolica var.grosseserrata were studied, and simulated herbivory experiments were carried out in order to evaluate leaf trait responses. Leaves ofQ. mongolica emerged simultaneously in spring and most were retained until autumn. Nitrogen concentration was highest when leaves first emerged and decreased rapidly with leaf age. Leaf mass per area (LMA) increased with leaf age. Herbivore attack was concentrated in the first 20 days after bud-break, which corresponded to the high nutritional value of the leaves for herbivores at this time. Simulated herbivory experiments indicated that LMA increased with artificial leaf damage, suggesting an increase in leaf toughness, and that nitrogen concentration decreased later in the season in comparison with intact leaves. As a result, herbivore attack following artificial leaf damage decreased with increasing initial leaf damage. However, leaf longevity was not affected by initial leaf damage. These responses were considered to be a strategy to disperse herbivory damage among leaves.     

Photosynthesis and resource distribution through plant canopies

Plant canopies are characterized by dramatic gradients of light between canopy top and bottom, and interactions between light, temperature and water vapour deficits. This review summarizes current knowledge of potentials and limitations of acclimation of foliage photosynthetic capacity (A(max)) and light-harvesting efficiency to complex environmental gradients within the canopies. Acclimation of A(max) to high light availability involves accumulation of rate-limiting photosynthetic proteins per unit leaf area as the result of increases in leaf thickness in broad-leaved species and volume: total area ratio and mesophyll thickness in species with complex geometry of leaf cross-section. Enhancement of light-harvesting efficiency in low light occurs through increased chlorophyll production per unit dry mass, greater leaf area per unit dry mass investment in leaves and shoot architectural modifications that improve leaf exposure and reduce within-shoot shading. All these acclimation responses vary among species, resulting in species-specific use efficiencies of low and high light. In fast-growing canopies and in evergreen species, where foliage developed and acclimated to a certain light environment becomes shaded by newly developing foliage, leaf senescence, age-dependent changes in cell wall characteristics and limited foliage re-acclimation capacity can constrain adjustment of older leaves to modified light availabilities. The review further demonstrates that leaves in different canopy positions respond differently to dynamic fluctuations in light availability and to multiple environmental stresses. Foliage acclimated to high irradiance respond more plastically to rapid changes in leaf light environment, and is more resistant to co-occurring heat and water stress. However, in higher light, co-occurring stresses can more strongly curb the efficiency of foliage photosynthetic machinery through reductions in internal diffusion conductance to CO(2). This review demonstrates strong foliage potential for acclimation to within-canopy environmental gradients, but also highlights complex constraints on acclimation and foliage functioning resulting from light x foliage age interactions, multiple environmental stresses, dynamic light fluctuations and species-specific leaf and shoot structural constraints.     

Leaf dynamics of seedlings of rain forest species in relation to canopy gaps

Summary Leaf dynamics of eight tropical rain forest species seedlings was studied in three environments: the shaded forest understorey, a small gap of ±50 m², and a large gap of ±500 m². Leaf production rate and leaf loss rate were enhanced in gaps, and a large gap resulted in larger increases than a small gap. For most species net leaf gain rate was larger in gaps, although this rate was not always largest in the large gap. Leaf loss decreased, and leaf survival percentages increased with increasing shade tolerance of species, indicating a slower leaf turnover for more shade tolerant species. Leaf area growth rate was only partly determined by net leaf gain rate. Ontogenetic effects on leaf size were also important, especially in the large gap. Species which possessed leaves with high specific leaf weight (SLW) showed lower leaf loss rates and higher leaf survival percentages than species with low SLW leaves. Leaf life span seemed to be related to leafcost per unit area. The relation of specific patterns in leaf production and leaf loss to the regeneration mode of the species is briefly discussed.     

Leaf structural and photosynthetic characteristics,and biomass allocation to foliage in relation to foliar nitrogen content and tree size in three Betula species

Young trees 0.03-1.7 m high of three coexisting Betula species were investigated in four sites of varying soil fertility, but all in full daylight, to separate nutrient and plant size controls on leaf dry mass per unit area (MA), light-saturated foliar photosynthetic electron transport rate (J) and the fraction of plant biomass in foliage (F(L)). Because the site effect was generally non-significant in the analyses of variance with foliar nitrogen content per unit dry mass (N(M)) as a covariate, N(M) was used as an explaining variable of leaf structural and physiological characteristics. Average leaf area (S) and dry mass per leaf scaled positively with N(M) and total tree height (H) in all species. Leaf dry mass per unit area also increased with increasing H, but decreased with increasing N(M), whereas the effects were species-specific. Increases in plant size led to a lower and increases in N(M) to a greater FL and total plant foliar area per unit plant biomass (LAR). Thus, the self-shading probably increased with increasing N(M) and decreased with increasing H. Nevertheless, the whole-plant average M(A), as well as M(A) values of topmost fully exposed leaves, correlated with N(M) and H in a similar manner, indicating that scaling of MA with N(M) and H did not necessarily result from the modified degree of within-plant shading. The rate of photosynthetic electron transport per unit dry mass (J(M)) scaled positively with N(M), but decreased with increasing H and M(A). Thus, increases in M(A) with tree height and decreasing nitrogen content not only resulted in a lower plant foliar area (LAR = F(L)/M(A)), but also led to lower physiological activity of unit foliar biomass. The leaf parameters (J(M), N(M) and M(A)) varied threefold, but the whole-plant characteristic FL varied 20-fold and LAR 30-fold, indicating that the biomass allocation was more plastically adjusted to different plant internal nitrogen contents and to tree height than the foliar variables. Our results demonstrate that: (1) tree height and N(M) may independently control foliar structure and physiology, and have an even greater impact on biomass allocation; and (2) the modified within-plant light availabilities alone do not explain the observed patterns. Although there were interspecific differences with respect to the statistical significance of the relationships, all species generally fit common regressions. However, these differences were consistent, and suggested that more competitive species with inherently larger growth rates also more plastically respond to N and H.     

A leaf gas exchange model that accounts for intra-canopy variability by considering leaf nitrogen content and local acclimation to radiation in grapevine (Vitis vinifera L.)

Understanding the distribution of gas exchange within a plant is a prerequisite for scaling up from leaves to canopies. We evaluated whether leaf traits were reliable predictors of the effects of leaf ageing and leaf irradiance on leaf photosynthetic capacity (V(cmax) , J(max) ) in field-grown vines (Vitis vinifera L). Simultaneously, we measured gas exchange, leaf mass per area (LMA) and nitrogen content (N(m) ) of leaves at different positions within the canopy and at different phenological stages. Daily mean leaf irradiance cumulated over 10 d (PPFD(10) ) was obtained by 3D modelling of the canopy structure. N(m) decreased over the season in parallel to leaf ageing while LMA was mainly affected by leaf position. PPFD(10) explained 66, 28 and 73% of the variation of LMA, N(m) and nitrogen content per area (N(a) ), respectively. Nitrogen content per unit area (N(a) = LMA × N(m) ) was the best predictor of the intra-canopy variability of leaf photosynthetic capacity. Finally, we developed a classical photosynthesis-stomatal conductance submodel and by introducing N(a) as an input, the model accurately simulated the daily pattern of gas exchange for leaves at different positions in the canopy and at different phenological stages during the season.     

Hydraulic conductance of leaves correlates with leaf lifespan: implications for lifetime carbon gain

Previous research suggests that the lifetime carbon gain of a leaf is constrained by a tradeoff between metabolism and longevity. The biophysical reasons underlying this tradeoff are not fully understood. We used a photosynthesis-leaf water balance model to evaluate biophysical constraints on carbon gain. Leaf hydraulic conductance (K(Leaf)), carbon isotope discrimination (Δ(13)C), leaf mass per unit area (LMA) and the driving force for water transport from stem to leaf (ΔΨ(Stem-Leaf)) were characterized for leaves spanning three orders of magnitude in surface area and two orders of magnitude in lifespan. We observed positive isometric scaling between K(Leaf) and leaf area but no relationship between Δ(13)C and leaf area. Leaf lifespan and LMA had minimal effect on K(Leaf) per unit leaf area, but a negative correlation exists among LMA, lifespan, and K(Leaf) per unit dry mass. During periods of leaf water loss, ΔΨ(Stem-Leaf) was relatively constant. We show for the first time that K(Leaf, mass), an index of the carbon cost associated with water use, is negatively correlated with lifespan. This highlights the importance of characterizing K(Leaf, mass) and suggests a tradeoff between resource investment in liquid phase processes and structural rigidity.     

Nitrogen reallocation and photosynthetic acclimation in response to partial shading in soybean plants

The first trifoliate of soybean was shaded when fully expanded, while the plant remained in high light; a situation representative for plants growing in a closed crop. Leaf mass and respiration rate per unit area declined sharply in the first few days upon shading and remained rather constant during the further 12 days of the shading treatment. Leaf nitrogen per unit area decreased gradually until the leaves were shed. Leaf senescence was enhanced by the shading treatment in contrast to control plants growing in low light. Shaded leaves on plants grown at low nutrient availability senesced earlier than shaded leaves on plants grown at high nutrient availability. The light saturated rate of photosynthesis decreased also gradually during the shading treatment, but somewhat faster than leaf N, whereas chlorophyll contents declined somewhat slower than leaf N.
Partitioning of N in the leaf over main photosynthetic functions was estimated from parameters derived from the response of photosynthesis to CO₂. It appeared that the N exported from the leaf was more at the expense of compounds that make up photosynthetic capacity than of those involved in photon absorption, resulting in a change in partitioning of N within the photosynthetic apparatus. Photosynthetic nitrogen use efficiency increased during the shading treatment, which was for the largest part due to the decrease in leaf N content, to some extent to the decrease in respiration rate and only for a small part to change in partitioning of N within the photosynthetic apparatus.     

Leaf traits in Chilean matorral: sclerophylly within,among, and beyond matorral,and its environmental determinants

Studies of leaf traits often focus on tradeoffs between growth and resource conservation, but little is known about variation in the mechanical traits that influence resource conservation. This study investigates how leaf mechanical traits vary across matorral vegetation in central Chile, how they correlate with environmental factors, and how these trends compare at a broader geographic scale. Leaf toughness, strength, stiffness, and associated traits were measured in five matorral types in central Chile, and relationships with soil N and P and climate variables were assessed. Trends with soil and climate were then analyzed across shrubland and woodland in Chile, Western Australia, and New Caledonia. Chilean species varied in leaf mechanics and associated traits, both within and among matorral types, with more species in sclerophyll matorral having strong, tough, and stiff leaves than in arid and littoral matorral. Overall, leaves with high leaf dry mass per area were stiffer, tougher, stronger, thicker, denser, with more fiber, lignin, phenolics and fiber per unit protein and less protein: tannin activity and N and P per mass, forming a broad sclerophylly syndrome. Mechanical traits of matorral species were not correlated with soil N or P, or predictably with climate variables, except flexural stiffness (EI_W) which correlated positively with annual reference evapotranspiration (ET₀). However, soil P made strong independent contributions to variation in leaf mechanics across shrublands and woodlands of Chile, Western Australia, and New Caledonia, either separately (strength) or together with ET₀ (toughness) explaining 46–90% of variation. Hence ET₀ was predictive of EI_W in Chilean matorral, whereas soil P was highly predictive of variation in leaf strength, and combined with ET₀ was highly predictive of toughness, at a broader geographic scale. The biological basis of these relationships, however, may be complex.     

The analysis of the causes of variability of the relationship between leaf dry mass and area in plants

The lamina dry mass: area ratio (LMA - Leaf Mass per Area) is a quite variable trait. Leaf dry mass consists of symplast mass (a set of all leaf protoplasts) and apoplast mass (a set of all cell walls in a leaf). The ratio between symplast and apoplast masses is positively related to any functional trait of leaf calculated per unit of dry mass. The value of this ratio is defined by cells size and their number per unit of leaf area, number of mesophyll cells layers and their differentiation between palisade and spongy ones, and also by density of cells packing. The LMA value is defined by leaf thickness and density. The extent and direction of variability in both leaf traits define the extent and direction of variability in LMA. Negative correlation between leaf thickness and density reduces the level of LMA variability. As a consequence of this correlation the following pattern emerges: the thinner a leaf, the denser it is. Changes in the traits that define the LMA value take place both within a species under the influence of environmental factors and between species that differ in leaf structure and functions. Light is the most powerful environmental factor that influences the LMA, increase in illumination leading to increase in LMA. This effect occurs during leaf growth at the expense of structural changes associated with the reduction of symplast/apoplast mass ratio. Under conditions of intense illumination, LMA may increase due to accumulation of starch. With regard to the majority of leaf functions, the mass of starch may be ascribed to apoplast. Starch accumulation in leaves is observed also under conditions of elevated CO2 concentration in the air. Under high illumination, however, LMA increases also due to increased apoplast contribution to leaf dry mass. Scarce mineral nutrition leads to LMA increase due to lowering of growth zones demands for phothosyntates and, therefore, to increase in starch content of leaves. High level of mineral nutrition during leaf growth period leads to LMA increase at the expense of mesophyll thickening where components of photosynthesis system are located. When additional environmental factors are involved, starch accumulation may be partly responsible for increase in LMA. LMA increase at the expense of starch accumulation, unlike that at the expense of mesophyll thickening, is accompanied by increased leaf density. Under conditions of water deficiency LMA increases, which in mature leaf may be caused by starch accumulation. LMA increase during leaf growth period under conditions of water deficiency is associated with decrease in the symplast/apoplast mass ratio.