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  • 1
    Electronic Resource
    Electronic Resource
    Springer
    Trees 14 (1999), S. 55-64 
    ISSN: 0931-1890
    Keywords: Key words Canopy ; Leaf angle distribution ; Shoot bifurcation ; Leaf area index ; Model
    Source: Springer Online Journal Archives 1860-2000
    Topics: Biology , Agriculture, Forestry, Horticulture, Fishery, Domestic Science, Nutrition
    Notes: 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.
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  • 2
    Electronic Resource
    Electronic Resource
    Oxford, UK : Blackwell Science Ltd
    Plant, cell & environment 28 (2005), S. 0 
    ISSN: 1365-3040
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology
    Notes: The present study was performed to investigate the adjustment of the rate parameters of the light and dark reactions of photosynthesis to the natural growth light in leaves of an overstorey species, Betula pendula Roth, a subcanopy species, Tilia cordata P. Mill., and a herb, Solidago virgaurea L., growing in a natural plant community in Järvselja, Estonia. Shoots were collected from the site and individual leaves were measured in a laboratory applying a standardized routine of kinetic gas exchange, Chl fluorescence and 820 nm transmittance measurements. These measurements enabled the calculations of the quantum yield of photosynthesis and rate constants of excitation capture by photochemical and non-photochemical quenchers, rate constant for P700+ reduction via the cytochrome b6f complex with and without photosynthetic control, actual maximum and potential (uncoupled) electron transport rate, stomatal and mesophyll resistances for CO2 transport, Km(CO2) and Vm of ribulose-bisphosphate carboxylase-oxygenase (Rubisco) in vivo. In parallel, N, Chl and Rubisco contents were measured from the same leaves. No adjustment toward higher quantum yield in shade compared with sun leaves was observed, although relatively more N was partitioned to the light-harvesting machinery in shade leaves (H. Eichelmann et al., 2004). The electron transport rate through the Cyt b6f complex was strongly down-regulated under saturating light compared with darkness, and this was observed under atmospheric, as well as saturating CO2 concentration. In vivo Vm measurements of Rubisco were lower than corresponding reported measurements in vitro, and the kcat per reaction site varied widely between leaves and growth sites. The correlation between Rubisco Vm and the photosystem I density was stronger than between Vm and the density of Rubisco active sites. The results showed that the capacity of the photosynthetic machinery decreases in shade-adjusted leaves, but it still remains in excess of the actual photosynthetic rate. The photosynthetic control systems that are targeted to adjust the photosynthetic rate to meet the plant's needs and to balance the partial reactions of photosynthesis, down-regulate partial processes of photosynthesis: excess harvested light is quenched non-photochemically; excess electron transport capacity of Cyt b6f is down-regulated by ΔpH-dependent photosynthetic control; Rubisco is synthesized in excess, and the number of activated Rubisco molecules is controlled by photosystem I-related processes. Consequently, the nitrogen contained in the components of the photosynthetic machinery is not used at full efficiency. The strong correlation between leaf nitrogen and photosynthetic performance is not due to the nitrogen requirements of the photosynthetic apparatus, but because a certain amount of energy must be captured through photosynthesis to maintain this nitrogen within a leaf.
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  • 3
    ISSN: 1365-3040
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology
    Notes: The present study was performed to investigate the adjustment of the constituents of the light and dark reactions of photosynthesis to the natural growth irradiance in the leaves of an overstorey species, Betula pendula Roth, a subcanopy species Tilia cordata P. Mill., and a herb Solidago virgaurea L. growing in a natural plant community in Järvselja, Estonia. Shoots were collected from the site and properties of individual leaves were measured in a laboratory, by applying a routine of kinetic gas exchange and optical measurements that revealed photosystem II (PSII), photosystem I (PSI), and cytochrome b6f densities per leaf area and the distribution of excitation (or chlorophyll, Chl) between the two photosystems. In parallel, N, Chl and ribulose-bisphosphate carboxylase-oxygenase (Rubisco) content was measured from the same leaves. The amount of N in photosynthetic proteins was calculated from the measured contents of the components of the photosynthetic machinery. Non-photosynthetic N was found as the residual of the budget. Growth in shade resulted in the decrease of leaf dry mass to a half of the DW in sun leaves in each species, but the total variation, from the top to the bottom of the canopy, was larger. Through the whole cross-section of the canopy, leaf dry weight (DW) and Rubisco content per area decreased by a factor of four, N content by a factor of three, but Chl content only by a factor of 1.7. PSII density decreased by a factor of 1.9, but PSI density by a factor of 3.2. The density of PSI adjusted to shade to a greater extent than the density of PSII. In shade, the distribution of N between the components of the photosynthetic machinery was shifted toward light-harvesting proteins at the expense of Rubisco. Non-photosynthetic N decreased the most substantially, from 54% in the sun leaves of B. pendula to 11% in the shade leaves of T. cordata. It is concluded that the redistribution of N toward light-harvesting Chl proteins in shade is not sufficient to keep the excitation rate of a PSII centre invariant. Contrary to PSII, the density of PSI – the photosystem that is in immediate contact with the carbon assimilation system – shade-adjusts almost proportionally with the latter, whereas its Chl antenna correspondingly increases. Even under N deficiency, a likely condition in the natural plant community, a substantial part of N is stored in non-photosynthetic proteins under abundant irradiation, but much less under limiting irradiation. At least in trees the general sequence of down-regulation due to shade adjustment is the following: (1) non-protein cell structures and non-photosynthetic proteins; (2) carbon assimilation proteins; (3) light reaction centre proteins, first PSI; and (4) chlorophyll-binding proteins.
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  • 4
    ISSN: 1365-3040
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology
    Notes: Responses of foliar light-saturated net assimilation rate (Amax), capacity for photosynthetic electron transport (Jmax) and mitochondrial respiration rate (Rd) to long-term canopy light and temperature environment were investigated in a temperate deciduous canopy composed of Populus tremula L. in the upper (17–28 m) and of Tilia cordata Mill. in the lower canopy layer (4–17 m). Climatic measurements indicated that seasonal average daily maximum air temperature (Tmax) was 5·5 °C (range 0·7–10·5 °C) higher in the top than in the bottom of the canopy, and strong positive correlations were observed between Tmax and seasonal average integrated quantum flux density (Qint), as well as between seasonal average daily mean temperature and Qint. Because of changes in leaf dry mass and nitrogen per unit area, Amax, Jmax, and Rd scaled positively with Qint in both species at a common leaf temperature (T). According to Jmax versus T response curves and dark chlorophyll fluorescence transients, photosynthetic electron transport was less heat resistant in P. tremula with optimum temperature of Jmax, Topt, of 33·5 ± 0·6 °C than in T. cordata with Topt of 40·7 ± 0·6 °C. This difference was suggested to manifest evolutionary adaptation of photosynthetic electron transport to cooler environments in P. tremula, the range of which extends farther north than that in T. cordata. Possibly because of acclimation to long-term canopy temperature environment, Topt was positively related to Qint in P. tremula, foliage of which was also exposed to higher irradiances and temperatures, but not in T. cordata, in the canopy of which quantum flux densities and temperatures were lower, and gradients in the environmental factors less pronounced. Parallel to changes in Topt, the activation energy for photosynthetic electron transport decreased with increasing Qint in P. tremula, indicating that Jmax of leaves acclimated to colder environment was more responsive to T in lower temperatures than that of high T acclimated leaves. Similar alterations in the activation energy for mitochondrial respiration rate were also observed, indicating that acclimation to temperature of mitochondrial and chloroplastic electron transport proceeds in a co-ordinated manner, and possibly involves long-term changes in membrane fluidity properties. We conclude that, because of correlations between temperature and light, the shapes of Jmax versus T, and Rd versus T response curves vary within tree canopies, and this needs to be taken account in modelling whole canopy photosynthesis.
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  • 5
    Electronic Resource
    Electronic Resource
    Oxford, UK : Blackwell Publishing Ltd
    Plant, cell & environment 18 (1995), S. 0 
    ISSN: 1365-3040
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology
    Notes: A simple analytical scheme, involving the distribution of nitrogen, to scale up photosynthesis from leaf to canopy is proposed. The scheme is based on the assumption that there are two pools of nitrogen in leaves: nitrogen in photosynthetic, degradable structures (Np) and nitrogen in non-photosynthetic and non-degradable structures (Ns). The rate of photon-saturated photosynthesis, Fm, is assumed to be proportional to Np and is distributed inside the canopy similarly to photon flux density (PFD). Prior assumptions of an optimum distribution of nitrogen are not a prerequisite. Calculations made with the scheme lead to development of the hypothesis that the canopy can be treated as a ‘big leaf’ on the time scales involved in acclimation of photosynthesis to PFD. Simulations using parameters for tree species with different requirements for PFD show that shade-tolerant species may have denser canopies than sun-demanding species because of smaller amounts of non-photosynthetic structural nitrogen and/or supporting tissue in their leaves.
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  • 6
    ISSN: 1365-3040
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology
    Notes: The observation of acclimation in leaf photosynthetic capacity to differences in growth irradiance has been widely used as support for a hypothesis that enables a simplification of some soil-vegetation-atmosphere transfer (SVAT) photosynthesis models. The acclimation hypothesis requires that relative leaf nitrogen concentration declines with relative irradiance from the top of a canopy to the bottom, in 1 : 1 proportion. In combination with a light transmission model it enables a simple estimate of the vertical profile in leaf nitrogen concentration (which is assumed to determine maximum carboxylation capacity), and in combination with estimates of the fraction of absorbed radiation it also leads to simple ‘big-leaf’ analytical solutions for canopy photosynthesis. We tested how forests deviate from this condition in five tree canopies, including four broadleaf stands, and one needle-leaf stand: a mixed-species tropical rain forest, oak (Quercus petraea (Matt.) Liebl), birch (Betula pendula Roth), beech (Fagus sylvatica L.) and Sitka spruce (Picea sitchensis (Bong.) Carr). Each canopy was studied when fully developed (mid-to-late summer for temperate stands). Irradiance (Q, µmol m−2 s−1) was measured for 20 d using quantum sensors placed throughout the vertical canopy profile. Measurements were made to obtain parameters from leaves adjacent to the radiation sensors: maximum carboxylation and electron transfer capacity (Va, Ja, µmol m−2 s−1), day respiration (Rda, µmol m−2 s−1), leaf nitrogen concentration (Nm, mg g−1) and leaf mass per unit area (La, g m−2).Relative to upper-canopy values, Va declined linearly in 1 : 1 proportion with Na. Relative Va also declined linearly with relative Q, but with a significant intercept at zero irradiance (P 〈 0·01). This intercept was strongly related to La of the lowest leaves in each canopy (P 〈 0·01, r2 = 0·98, n= 5). For each canopy, daily lnQ was also linearly related with lnVa(P 〈 0·05), and the intercept was correlated with the value for photosynthetic capacity per unit nitrogen (PUN: Va/Na, µmol g−1 s−1) of the lowest leaves in each canopy (P 〈 0·05). Va was linearly related with La and Na(P 〈 0·01), but the slope of the Va : Na relationship varied widely among sites. Hence, whilst there was a unique Va : Na ratio in each stand, acclimation in Va to Q varied predictably with La of the lowest leaves in each canopy. The specific leaf area, Lm(cm2 g−1), of the canopy-bottom foliage was also found to predict carboxylation capacity (expressed on a mass basis; Vm, µmol g−1 s−1) at all sites (P 〈 0·01). These results invalidate the hypothesis of full acclimation to irradiance, but suggest that La and Lm of the most light-limited leaves in a canopy are widely applicable indicators of the distribution of photosynthetic capacity with height in forests.
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  • 7
    ISSN: 1365-3040
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology
    Notes: The present study was undertaken to test for the hypothesis that the rate of development in the capacity for photosynthetic electron transport per unit area (Jmax;A), and maximum carboxylase activity of Rubisco (Vcmax;A) is proportional to average integrated daily quantum flux density (Qint) in a mixed deciduous forest dominated by the shade-intolerant species Populus tremula L., and the shade-tolerant species Tilia cordata Mill. We distinguished between the age-dependent changes in net assimilation rates due to modifications in leaf dry mass per unit area (MA), foliar nitrogen content per unit dry mass (NM), and fractional partitioning of foliar nitrogen in the proteins of photosynthetic electron transport (FB), Rubisco (FR) and in light-harvesting chlorophyll-protein complexes (Vcmax;A ∝ MANMFR; Jmax;A ∝ MANMFB). In both species, increases in Jmax;A and Vcmax;A during leaf development were primarily determined by nitrogen allocation to growing leaves, increases in leaf nitrogen partitioning in photosynthetic machinery, and increases in MA. Canopy differences in the rate of development of leaf photosynthetic capacity were mainly controlled by the rate of change in MA. There was only small within-canopy variation in the initial rate of biomass accumulation per unit Qint (slope of MA versus leaf age relationship per unit Qint), suggesting that canopy differences in the rate of development of Jmax;A and Vcmax;A are directly proportional to Qint. Nevertheless, MA, nitrogen, Jmax;A and Vcmax;A of mature leaves were not proportional to Qint because of a finite MA in leaves immediately after bud-burst (light-independent component of MA). MA, leaf chlorophyll contents and chlorophyll : N ratio of mature leaves were best correlated with the integrated average quantum flux density during leaf development, suggesting that foliar photosynthetic apparatus, once developed, is not affected by day-to-day fluctuations in Qint. However, for the upper canopy leaves of P. tremula and for the entire canopy of T. cordata, there was a continuous decline in N contents per unit dry mass in mature non-senescent leaves on the order of 15–20% for a change of leaf age from 40 to 120 d, possibly manifesting nitrogen reallocation to bud formation. The decline in N contents led to similar decreases in leaf photosynthetic capacity and foliar chlorophyll contents. These data demonstrate that light-dependent variation in the rate of developmental changes in MA determines canopy differences in photosynthetic capacity, whereas foliar photosynthetic apparatus is essentially constant in fully developed leaves.
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  • 8
    ISSN: 1365-3040
    Source: Blackwell Publishing Journal Backfiles 1879-2005
    Topics: Biology
    Notes: To test the hypothesis that in temperate deciduous trees acclimation to potentially damaging high irradiances occurs via long-term adjustments in foliar photosynthetic capacity, and short-term changes in xanthophyll cycle pool size in response to weather fluctuations, nitrogen concentration and pigment composition were examined along a canopy light gradient in three species –Betula pendula, Populus tremula and Tilia cordata (from most shade intolerant to tolerant), and foliage photosynthetic potentials in P. tremula and T. cordata. Integrated quantum flux density (Qi) incident on leaves was estimated with a method combining hemispherical photography and light measurements with quantum sensors made over the growing season. Long- and short-term light indices – average total seasonal daily integrated quantum flux density (Ts, mol m–2 d–1) and that of the 3 d preceding foliage sampling (T3d) – were calculated for each sampled leaf. In addition to total integrated quantum flux density, the part of Qi attributable to direct flux was also computed. Strong linear relationships between the capacity for photosynthetic electron transport per area (Jamax), estimated from in situ measurements of effective quantum yield of photosystem II (PS II), and Qi averaged over the season and over the preceding 3 d were found for all studied species. However, the major determinant of Jamax, the product of electron transport capacity per leaf dry mass (Jmmax) and leaf dry mass per area (MA), was MA rather than Jmmax, which was relatively constant along the light gradient. There was evidence that Jamax is more tightly related to Ts, which characterizes the light climate during foliar development, than to short-term integrated light, possibly because there is little flexibility in adjustments in MA after the completion of foliar growth. Leaf chlorophyll concentrations and the investment of leaf nitrogen in chlorophyll (Chl/N) were negatively related to Qi– an investment pattern which improves light harvesting in low light. Xanthophyll cycle pool size (VAZ, violaxanthin + antheraxanthin + zeaxanthin) either expressed per unit chlorophyll (VAZ/Chl) or as a fraction of total carotenoids (VAZ/Car) increased with increasing Qi in all species. However, contrary to Jamax, it tended to correlate more strongly with short-term than with long-term average integrated light. There were few interspecific differences in Jamax, Chl/N, VAZ/Chl and VAZ/Car when the variability in light level incident to the leaves was accounted for, indicating that the foliage of both shade-intolerant and -tolerant temperate tree species possesses considerable phenotypic flexibility. Collectively these results support the view that rapid adjustment of the xanthophyll cycle pool size provides an important means for acclimation to light fluctuations in a time scale of days, during which the potential for photosynthetic quenching of excitation energy is not likely to change appreciably.
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  • 9
    Publication Date: 2013-02-21
    Description: We investigated changes in chlorophyll a fluorescence from alternate leaf surfaces to assess the intraleaf light acclimation patterns in combination with natural variations in radiation, leaf angles, leaf mass per area (LMA), chlorophyll content (Chl) and leaf optical parameters. Measurements were conducted on bottom- and top-layer leaves of Tilia cordata Mill. (a shade-tolerant sub-canopy species, sampled at heights of 11 and 16 m) and Populus tremula L. (a light-demanding upper canopy species, sampled at canopy heights of 19 and 26 m). The upper canopy species P. tremula had a six times higher PSII quantum yield ( II ) and ratio of open reaction centres (qP), and a two times higher LMA than T. cordata . These species-specific differences were also present when the leaves of both species were in similar light conditions. Leaf adaxial/abaxial fluorescence ratio was significantly larger in the case of more horizontal leaves. Populus tremula (more vertical leaves), had smaller differences in fluorescence parameters between alternate leaf sides compared with T. cordata (more horizontal leaves). However, optical properties on alternate leaf sides showed a larger difference for P. tremula . Intraspecifically, the measured optical parameters were better correlated with LMA than with leaf Chl. Species-specific differences in leaf anatomy appear to enhance the photosynthetic potential of leaf biochemistry by decreasing the interception of excess light in P. tremula and increasing the light absorptance in T. cordata . Our results indicate that intraleaf light absorption gradient, described here as leaf adaxial/abaxial side ratio of chlorophyll a fluorescence, varies significantly with changes in leaf light environment in a multi-layer multi-species tree canopy. However, this variation cannot be described merely as a simple function of radiation, leaf angle, Chl or LMA, and species-specific differences in light acclimation strategies should also be considered.
    Print ISSN: 0829-318X
    Electronic ISSN: 1758-4469
    Topics: Agriculture, Forestry, Horticulture, Fishery, Domestic Science, Nutrition
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  • 10
    Publication Date: 1996-04-01
    Description: Current projections indicate steady increases in both trophospheric ozone and carbon dioxide well into the next century with concurrent increases in plant stress. Because information about effects of these interacting stresses on forest trees is limited, we have conducted ozone and carbon dioxide experiments using ozone-tolerant and ozone-sensitive trembling aspen (Populustremuloides Michx.) clones (clones 216 and 259, respectively). Aspen plants were grown either in pots (square-wave study) or in the ground (episodic study) in open-top chambers. Plants in the square-wave study were exposed for a single growing season to charcoal-filtered air (CF) or to CF plus elevated carbon dioxide (CO2), ozone (O3), or O3 plus CO2 (O3 + CO2). Plants in the episodic study were exposed for three growing seasons to CF, twice simulated ambient (2x) O3 (2x O3), or 2x O3 plus CO, (2x O3 + CO2). Photosynthetic measurements were made either in the open-top chambers at treatment CO2 concentrations or in controlled-environment cuvettes with various CO2 concentrations, producing assimilation versus intercellular CO2 concentration (A/Ci) curves. Ozone decreased photosynthetic rate and stomatal conductance and accelerated leaf senescence. Elevated CO2 increased photosynthetic rate and decreased stomatal conductance when measured at treatment CO2 concentrations, and exacerbated the negative effect of O3 on photosynthesis. For example, for clone 259, photosynthesis decreased 9% for the O3 treatment compared with the CF treatment, but decreased 24% for the O3 + CO2 treatment compared with the CF treatment. Similar decreases for clone 216 of 2% and 6% for O3 and O3 + CO2, respectively, were not significant. A/Ci curves showed that O3 decreased carboxylation efficiency and maximum photosynthetic rate and that photosynthetic inhibition in response to O3 was greater with elevated CO2. The simultaneous declines in all factors of photosynthetic gas exchange measurements suggest that the equilibrium between stomatal conductance, carboxylation, and light harvesting systems was not disrupted by O3 and O3× CO2 interactions. Carbon dioxide did not ameliorate the detrimental effects of O3 on the leaf photosynthetic apparatus. In fact, the O3-tolerant clone appeared more sensitive to O3 with elevated CO2.
    Print ISSN: 0045-5067
    Electronic ISSN: 1208-6037
    Topics: Agriculture, Forestry, Horticulture, Fishery, Domestic Science, Nutrition
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