Olevi Kull

Acclimation of photosynthesis in canopies: models and limitations

Abstract. Within a time-scale of several days photosynthesis can acclimate to light by variation in the capacity for photosynthesis with depth in a canopy or by variation in the stoichiometry of photosynthetic components at each position within the canopy. The changes in leaf photosynthetic capacity are usually related to and expressed as changes in leaf nitrogen content. However, photosynthetic capacity and leaf nitrogen never match exactly the photon flux density (PFD) gradient within a canopy. As a result, photosynthetic light use efficiency, i.e. photosynthetic performance per incident PFD, increases considerably from the top of the canopy to the lower shaded part. Many of existing optimisation models fail to express the actual pattern of nitrogen or photosynthetic capacity distribution within a canopy. This failure occurs because these optimisation models do not consider that the quantitative aspect of photosynthesis acclimation is a whole plant phenomenon. Although turnover models, which describe the distribution of the photosynthetic apparatus within a canopy as a dynamic equilibrium between breakdown and regeneration of apparatus with respect to nitrogen availability, photosynthetic rate and export of carbohydrates, produce realistic results, these models require confirmation. The mechanism responsible for changes in the relative share of light-harvesting apparatus as acclimation to irradiance remains unknown. Ability of the photosynthetic apparatus to balance properly the light harvesting capacity with electron transport and biochemical capacities is limited. As a result of this fundamental limitation, photosynthetic light use efficiency always increases with increasing thickness of the photosynthetic apparatus.

Variability in Leaf Morphology and Chemical Composition as a Function of Canopy Light Environment in Coexisting Deciduous Trees

Morphology, chemical composition, and photosynthetic capacity of leaf laminas were investigated in Populus tremula L. and Tilia cordata Mill. along a canopy light gradient. Variables determining the thickness of boundary layer for heat and water exchange at a given wind speed—effective leaf width (Ww) and length (Wd)—scaled positively with daily integrated quantum flux density averaged over the season (Qint, mol m−2 d−1) in T. cordata, but Wd decreased and Ww was constant with increasing Qint in P. tremula, bringing about a moderately improved capacity for convective cooling at greater irradiances in the latter species. Foliar stable carbon isotope discrimination (Δ) decreased with increasing Qint, demonstrating that, possibly because of more severe foliar water stress, leaves operated at a lower intercellular CO2 concentration in the upper canopy. Further analysis of foliar characteristics provided additional evidence of the interaction between water stress and Qint. Leaf dry matter content and both components of lamina dry mass per area (MA)—lamina thickness and density (dry mass per unit volume, ρB)—increased with increasing Qint in both species. The ρB and lamina dry matter content were also positively related to lamina carbon concentration, variability in which along the canopy was related to changes in carbon‐rich lignin concentration. Since both increases in lamina density and lignin concentration improve leaf tolerance of low‐water potentials, these foliar modifications were interpreted as indicative of acclimation to enhanced water limitations in high light. For the whole material, foliar nitrogen concentrations decreased with increasing ρB, suggesting that an improvement of foliar mechanical strength may result in declining foliar assimilative potential. However, foliar photosynthetic electron transport capacity per unit area increased with increasing ρB, possibly because increases in ρB with light are not only attributable to greater cell wall lignification but also to denser packing of leaf cells, in particular, in fractional increases in palisade tissues with Qint. Because of a positive scaling of leaf thickness and density with total tree height, MA was greater in taller trees of T. cordata, foliage of which also had lower Δ and was likely to function with less open stomata. In summary, we conclude that leaf water stress, which scales with both Qint and total tree height, is a major factor altering foliage structure and assimilative capacity.

Light distribution and foliage structure in an oak canopy

Abstract Leaf angle distribution and shoot bifurcation ratio were measured and related to photon flux density (PFD) distribution in an oak canopy. Leaf angle distribution deviated substantially from random and changed markedly throughout the canopy. The observed leaf angle distribution was described by an ellipsoidal function with the single parameter of the distribution, x, changing from 1.6 at the top of the canopy to 3.2 in the lowest branches. In vertically homogeneous canopies, the extinction coefficient for diffuse radiation is expected to decline with increasing leaf area index (LAI). However, in the canopy studied here, the leaf angle distribution changed with height such that the effective extinction coefficient remained constant. Both shoot bifurcation ratio and leaf number per shoot declined with decreasing PFD inside the canopy. Based on these observed relationships, a simple canopy growth model that assumes horizontal homogeneity of the canopy was constructed. Calculations showed that a steady state, when growth in the upper of the canopy is in equilibrium with decline of lower canopy, the total canopy LAI should equal to 4.3. This predicted value of equilibrium LAI is larger than that estimated from measurements of PFD transmission (LAI=3.3), but smaller than that directly determined by litter collection (LAI=6.2 in 1998). Possible reasons for these discrepancies are discussed.

Net photosynthetic response to light intensity of shoots from different crown positions and age in picea abies (L.) karst

Carbon dioxide uptake response to light intensity of detached shoots of open‐ and forest‐grown Norway spruces was investigated in controlled standard conditions. The initial slope of the CO2 uptake light response curve, calculated on leaf area basis and the internal conductance, calculated on leaf weight basis were almost constant for shoots from different positions in the forest canopy. In open‐grown trees the maximum photosynthesis and internal conductance (on leaf weight basis) decreased in relation to the tree age, so that in the 8‐year‐old tree it was about two times of that in the 66‐year‐old tree. These results suggest that 1) the enhanced efficiency of “shade” shoots is caused by morphologic adaptation, i.e. the more sparsely packed photosynthetic apparatus in needles and shoots, and 2) adaptational possibilities of Norway spruce photosynthetic apparatus, its ecological plasticity diminish during tree ontogenesis.

Apparent Controls on Leaf Conductance by Soil Water Availability and via Light‐Acclimation of Foliage Structural and Physiological Properties in a Mixed Deciduous, Temperate Forest

Controls on leaf stomatal conductances imposed by soil water availability and foliage acclimation to long‐term integrated irradiance were studied in a natural mixed deciduous stand composed of shade‐intolerant Populus tremula L. and shade‐tolerant Tilia cordata Mill. Positive relationships between maximum stomatal conductance and seasonal integrated average daily quantum flux density ( \documentclass{aastex} \usepackage{amsbsy} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{bm} \usepackage{mathrsfs} \usepackage{pifont} \usepackage{stmaryrd} \usepackage{textcomp} \usepackage{portland,xspace} \usepackage{amsmath,amsxtra} \usepackage[OT2,OT1]{fontenc} \newcommand\cyr{ \renewcommand\rmdefault{wncyr} \renewcommand\sfdefault{wncyss} \renewcommand\encodingdefault{OT2} \normalfont \selectfont} \DeclareTextFontCommand{\textcyr}{\cyr} \pagestyle{empty} \DeclareMathSizes{10}{9}{7}{6} \begin{document} \landscape

Photosynthetic acclimation to light in woody and herbaceous species: a comparison of leaf structure, pigment content and chlorophyll fluorescence characteristics measured in the field

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Elevated CO2 response of photosynthesis depends on ozone concentration in aspen.

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