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1.
以1-2年生北加州黑核桃为试材,建立了具有较高分辨能力的植株群体结构、光分布模型和冠层光合作用模型.将植株冠层按叶面积指数划分为若干层次。上下层之间水平面上太阳辐照度按Monsi&Saeki所提出的指数递减规律分布.冠层内太阳散射光的消光系数由冠层结构决定,而直射光的消光系数则决定于冠层结构与太阳在天空的位置.在同一层次。将叶片的叶倾角划分为6个等级。将叶片的水平位置划分为8个方位.设同一层次中水平面上的太阳辐照度相同。某一方位角和叶倾角的叶面的直接辐射由太阳视运动方程决定.以此为基础,分别计算“光斑区”和“遮荫区”内叶片的光合速率,并通过数值积分计算整个冠层的光合速率及光合日总量.用田间实测资料验证了冠层内太阳辐射分布模型和冠层光合作用模型.敏感性试验分析表明。模型对环境因子和生物学因素有良好的响应.  相似文献   

2.
Summary A hypothesis that a dense stand should develop a less uniform distribution of leaf nitrogen through the canopy than an open stand to increase total canopy photosynthesis was tested with experimentally established stands of Lysimachia vulgaris L. The effect of stand density on spatial variation of photon flux density, leaf nitrogen and specific leaf weight within the canopy was examined. Stand density had little effect on the value of the light extinction coefficient, but strongly affected the distribution of leaf nitrogen per unit area within a canopy. The open stand had more uniform distribution of leaf nitrogen than the dense stand. However, different light climates between stands explained only part of the variation of leaf nitrogen in the canopy. The specific leaf weight in the canopy increased with increasing relative photon flux density and with decreasing nitrogen concentration.  相似文献   

3.
Seasonal changes and yearly gross canopy photosynthetic production were estimated for an 18 year old Japanese larch (Larix leptolepis) forest between 1982 and 1984. A canopy photosynthesis model was applied for the estimation, which took into account the effect of light interception by the non-photosynthetic organs. Seasonal changes in photosynthetic ability, amount of canopy leaf area and light environment within the canopy were also taken into account. Amount of leaf area was estimated by the leaf area growth of a single leaf. The change of light environment within the canopy during the growing season was estimated with a light penetration model and the leaf increment within the canopy. Canopy respiration and surplus production were calculated as seasonal and yearly values for the three years studied. Mean yearly estimates of canopy photosynthesis, canopy respiration and surplus production were 37, 13 and 23 tCO2 ha−1 year−1, respectively. Vertical trend, seasonal changes and yearly values of the estimates were analyzed in relation to environmental and stand factors.  相似文献   

4.
Development of the Monsi-Saeki theory on canopy structure and function   总被引:11,自引:0,他引:11  
Hirose T 《Annals of botany》2005,95(3):483-494
BACKGROUND AND AIMS: Monsi and Saeki (1953) published the first mathematical model of canopy photosynthesis that was based on the light attenuation within a canopy and a light response of leaf photosynthesis. This paper reviews the evolution and development of their theory. SCOPE: Monsi and Saeki showed that under full light conditions, canopy photosynthesis is maximized at a high leaf area index (LAI, total leaf area per unit ground area) with vertically inclined leaves, while under low light conditions, it is at a low LAI with horizontal leaves. They suggested that actual plants develop a stand structure to maximize canopy photosynthesis. Combination of the Monsi-Saeki model with the cost-benefit hypothesis in resource use led to a new canopy photosynthesis model, where leaf nitrogen distribution and associated photosynthetic capacity were taken into account. The gradient of leaf nitrogen in a canopy was shown to be a direct response to the gradient of light. This response enables plants to use light and nitrogen efficiently, two resources whose supply is limited in the natural environment. CONCLUSION: The canopy photosynthesis model stimulated studies to scale-up from chloroplast biochemistry to canopy carbon gain and to analyse the resource-use strategy of species and individuals growing at different light and nitrogen availabilities. Canopy photosynthesis models are useful to analyse the size structure of populations in plant communities and to predict the structure and function of future terrestrial ecosystems.  相似文献   

5.
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.  相似文献   

6.
The importance of light acquisition and utilization by individuals in intraspecific competition was evaluated by determining growth and photosynthesis of individual plants in a dense monospecific stand of an annual, Xanthium canadense. Photosynthesis of individual plants in the stand was calculated using a canopy photosynthesis model in which leaf photosynthesis was assumed to be function of leaf nitrogen content and light availability. The estimated photosynthetic rates of individuals were strongly correlated with the measured growth rates. Photosynthetic rates per unit aboveground mass (RPR, relative photosynthetic rate) increased with increasing aboveground mass, suggesting asymmetric (one-sided) competition in the stand. However, larger individuals had similar RPRs, suggesting symmetric (two-sided) competition. These results were consistent with the observation that size inequality over the whole stand increased with growth, but it remained stable among the larger individuals. The RPR of an individual was calculated as the product of absorbed photon flux per unit aboveground mass (Φmass) and light use efficiency (LUE, photosynthesis per unit absorbed photon flux). Φmass indicates the efficiency of light acquisition, and was higher in larger individuals in the stand, while LUE was highest in individuals with intermediate aboveground mass. LUE depends on leaf nitrogen content. At an early stage, leaf nitrogen contents of smaller individuals were similar to those that maximize LUE. Light availability to smaller individuals decreased as they grew, while their nitrogen contents did not change markedly, which decreased their LUE. We concluded that asymmetric competition among individuals in the stand resulted mainly from lower efficiencies in both light acquisition and light use by smaller individuals. Received: 31 January 1998 / Accepted: 12 November 1998  相似文献   

7.
8.
Summary Seasonal changes in the light and temperature dependence of photosynthesis were investigated in field grown plants of Mercurialis perennis and Geum urbanum. In both species changes in photosynthetic capacity were closely related to the development of the overstorey canopy. In G. urbanum there was a marked shift in the temperature dependence of photosynthesis through the season whereas no such pattern was found in M. perennis. Model predictions of field rates of photosynthesis were made using the measurements of light and temperature dependence in the laboratory and validated against field observations. Long term continuous records of light and temperature in the field were used in conjunction with the model to make predictions of carbon acquisition in shoots of the two species through the season. These calculations indicated that G. urbanum was able to take advantage of high light levels just prior to canopy closure through a combination of high photosynthetic capacity, the ability to maintain photosynthesis at relatively low temperatures and the presence of overwintering leaves. In M. perennis leaf development was early enough to utilise the high spring light period. After canopy closure M. perennis maintained a higher average rate of CO2 flux due to a combination of high apparent quantum efficiency and low rates of respiration.  相似文献   

9.
In order to parametrize a leaf submodel of a canopy level gas-exchange model, a series of photosynthesis and stomatal conductance measurements were made on leaves of white oak (Quercus alba L.) and red maple (Acer rubrum L.) in a mature deciduous forest near Oak Ridge, TN. Gas-exchange characteristics of sun leaves growing at the top of a 30 m canopy and of shade leaves growing at a depth of 3–4 m from the top of the canopy were determined. Measured rates of net photosynthesis at a leaf temperature of 30°C and saturating photosynthetic photon flux density, expressed on a leaf area basis, were significantly lower (P = 0.01; n = 8) in shade leaves (7.9μmol m?2 s?1) than in sun leaves (11–5μmol m?2 s?1). Specific leaf area increased significantly with depth in the canopy, and when photosynthesis rates were expressed on a dry mass basis, they were not significantly different for shade and sun leaves. The percentage leaf nitrogen did not vary significantly with height in the canopy; thus, rates expressed on a per unit nitrogen basis were also not significantly different in shade and sun leaves. A widely used model integrating photosynthesis and stomatal conductance was parametrized independently for sun and shade leaves, enabling us to model successfully diurnal variations in photosynthesis and evapotranspiration of both classes of leaves. Key photosynthesis model parameters were found to scale with leaf nitrogen levels. The leaf model parametrizations were then incorporated into a canopy-scale gas-exchange model that is discussed and tested in a companion paper (Baldocchi & Harley 1995, Plant, Cell and Environment 18, 1157–1173).  相似文献   

10.
Photosynthetic capacity was measured on detached leaves sampled in a canopy of Solidago altissima L. Non-rectangular hyperbola fitted the light response curve of photosynthesis and significant correlations were observed between leaf nitrogen per unit area and four parameters which characterize the light-response curve. Using regressions of the parameters on leaf nitrogen, a model of leaf photosynthesis was constructed which gave the relationships between leaf nitrogen, photon flux density (PFD) and photosynthesis. Curvilinear relations were obtained between leaf nitrogen and photosynthetic rate on both an instantaneous and a daily basis. Nitrogen use efficiency (NUE, photosynthesis per unit leaf nitrogen) was calculated against leaf nitrogen under varying PFDs. The optimum nitrogen content per unit leaf area that maximizes NUE shifted to higher values with increasing PFD. Field measurements of PFD showed high positive correlations between the distribution of leaf nitrogen in the canopy and relative PFD. The predicted optimum leaf nitrogen content for each level in the canopy, to achieve maximized NUE during a clear day, was close to the actual nitrogen distribution as found through sampling.  相似文献   

11.
The rates of canopy and individual leaf photosynthesis, ratesof growth of shoots and roots, and the extinction coefficientfor light of eight temperate forage grasses were determinedin the field during early autumn. Canopy gross photosynthesiswas calculated as net photosynthesis plus dark respiration adjustedfor temperature using a Q10 = 2. The relationships between canopygross photosynthesis and light intensity were hyperbolic, andthe initial slopes of these curves indicated that light wasbeing utilized efficiently at low light intensities. The initialslope depended on the distribution of light in the canopy andthe quantum efficiency of the individual leaves. The maximumrate of canopy gross photosynthesis reflected the maximum rateof individual leaf photosynthesis. Although the maximum rateof canopy gross photosynthesis was correlated with crop growthrate, there was no significant relationship between daily grossphotosynthesis and crop growth rate. Indeed, daily gross photosynthesisvaried by only 22 per cent, whereas the daily growth of shootsand roots varied by 120 per cent. This poor correlation is influencedby a negative correlation (P < 0.01) between the maximumrate of canopy gross photosynthesis and the initial slope ofthe curve relating canopy gross photosynthesis and light intensity.Difficulties in the interpretation of measurements of dark respirationappeared to confound attempts to relate daily net photosynthesisto crop growth rate, individual leaf photosynthesis, and theextinction coefficient for light.  相似文献   

12.
BACKGROUND AND AIMS: Light extinction through crowns of canopy trees determines light availability at lower levels within forests. The goal of this paper is the exploration of foliage distribution and light extinction in crowns of five canopy tree species in relation to their shoot architecture, leaf traits (mean leaf angle, life span, photosynthetic characteristics) and successional status (from pioneers to persistent). METHODS: Light extinction was examined at three hierarchical levels of foliage organization, the whole crown, the outermost canopy and the individual shoots, in a tropical moist forest with direct canopy access with a tower crane. Photon flux density and cumulative leaf area index (LAI) were measured at intervals of 0.25-1 m along multiple vertical transects through three to five mature tree crowns of each species to estimate light extinction coefficients (K). RESULTS: Cecropia longipes, a pioneer species with the shortest leaf life span, had crown LAI <0.5. Among the remaining four species, crown LAI ranged from 2 to 8, and species with orthotropic terminal shoots exhibited lower light extinction coefficients (0.35) than those with plagiotropic shoots (0.53-0.80). Within each type, later successional species exhibited greater maximum LAI and total light extinction. A dense layer of leaves at the outermost crown of a late successional species resulted in an average light extinction of 61% within 0.5 m from the surface. In late successional species, leaf position within individual shoots does not predict the light availability at the individual leaf surface, which may explain their slow decline of photosynthetic capacity with leaf age and weak differentiation of sun and shade leaves. CONCLUSION: Later-successional tree crowns, especially those with orthotropic branches, exhibit lower light extinction coefficients, but greater total LAI and total light extinction, which contribute to their efficient use of light and competitive dominance.  相似文献   

13.
  • 1 The ‘big‐leaf’ approach to calculating the carbon balance of plant canopies is evaluated for inclusion in the ETEMA model framework. This approach assumes that canopy carbon fluxes have the same relative responses to the environment as any single leaf, and that the scaling from leaf to canopy is therefore linear.
  • 2 A series of model simulations was performed with two models of leaf photosynthesis, three distributions of canopy nitrogen, and two levels of canopy radiation detail. Leaf‐ and canopy‐level responses to light and nitrogen, both as instantaneous rates and daily integrals, are presented.
  • 3 Observed leaf nitrogen contents of unshaded leaves are over 40% lower than the big‐leaf approach requires. Scaling from these leaves to the canopy using the big‐leaf approach may underestimate canopy photosynthesis by ~20%. A leaf photosynthesis model that treats within‐leaf light extinction displays characteristics that contradict the big‐leaf theory. Observed distributions of canopy nitrogen are closer to those required to optimize this model than the homogeneous model used in the big‐leaf approach.
  • 4 It is theoretically consistent to use the big‐leaf approach with the homogeneous photosynthesis model to estimate canopy carbon fluxes if canopy nitrogen and leaf area are known and if the distribution of nitrogen is assumed optimal. However, real nitrogen profiles are not optimal for this photosynthesis model, and caution is necessary in using the big‐leaf approach to scale satellite estimates of leaf physiology to canopies. Accurate prediction of canopy carbon fluxes requires canopy nitrogen, leaf area, declining nitrogen with canopy depth, the heterogeneous model of leaf photosynthesis and the separation of sunlit and shaded leaves. The exact nitrogen profile is not critical, but realistic distributions can be predicted using a simple model of canopy nitrogen allocation.
  相似文献   

14.
Summary We studied the effects of density on the dynamics of seedling growth and canopy microclimate within experimental stands composed of Datura ferox L. seedlings grown in individual pots. Interception of photosynthetically active radiation (PAR) by seedlings was evaluated either indirectly, by measuring leaf area, proportion of leaf area shaded by neighbouring individuals and laminar orientation with respect to sunlight, or directly, by measuring PAR at individual leaves at their natural angle of display. An integrating cylinder, with a geometry approximating that of a stem, was used within the canopies to measure the red:far-red (R:FR) ratio of the light flux from all compass points parallel to the soil surface. Seedlings responded rapidly (i.e. 1–2 weeks) to increased density by producing longer internodes and partitioning more dry matter to stems relative to leaves. These responses were observed before either PAR interception of growth rate were reduced by the presence of neighbours. Conversely, morphogenetic adjustment was preceded by a consistent effect of plant density on the R:FR ratio of the light received by the integrating cylinder. Air and soil temperature were not affected by density in these experiments. Differences in wind velocity within the canopy associated with plant density were avoided by the experimental procedure. The results support the idea that the drop in R:FR ratio of the light flux parallel to the ground — e.g. reflected sunlight — is an early signal that allows rapid adjustment of plant form to changes in canopy structure.  相似文献   

15.
孙谷畴   《广西植物》1987,(3):239-243
亚热带季雨林林下阴生植物罗伞(Ardisia quinquegona)叶片的气体交换速率(PN.μmol.m~(-2),s~(-1))随光强(PFD,μmol,m~(-2),s~(-1))增高而增大。在光强低于80μmol,m~(-2),s~(-1),PN=29.21PFD×10~(-3)+0.36。在光强150μmol,m~(-2),s~(-1)对出现气体交换的光饱和现象。在低光强下,气孔传导率(G,m mol,m~(-2),s~(-1)与光强(m mol,m~(-2),s~(-1)的关系为G=265.6 PFD+4.6。在低光强下。开阔地的阳生灌木桃金娘(Rhodmyrtus tomentosa)的气体交换速率和气孔传导率与光强关系曲线的直线部分斜率皆较罗伞的低,在红光上,罗伞叶片气体交换速率(μmol,m~(-2),s~(-1)与光强(μmol,m~(-2),s~(-1)的关系为PN=32.4 PFD×10~(-3)-0.04。气孔传导率(m mol,m~(-2),s~(-1)与光强(m mol,m~(-2),s~(-1)的关系为G=339.08 PFD+7.37。同时气体交换速率的饱和红光光强亦较白光的高。在蓝光光强低时,气体交换速率(μmol,m~(-2),s~(-1))与光强(μmol,m~(-2),s~(-1))的关系为PN=13.54 PFD×10~(-3)—0.17,而气孔传导率(m mol,m~(-2),s~(-1))与光强(mμmol,m~(-2),s~(-1))的关系为G=80.5 PFD+4.35。在低的蓝光下,体交换速率和气孔传导率与光强关系曲线的直线部分斜率显著较在白光和红光下的低。罗伞叶片气体交换对红光的反应敏感。  相似文献   

16.
Most models for canopy photosynthesis require a large number of parameters as input which have to be determined by means of direct measurements. Such measurements are usually expensive, time consuming and destructive. The objective of the present study was, therefore, to develop a simple but accurate canopy photosynthesis model based on a minimum number of parameters that can be determined non-destructively. The results from previous studies were used to derive an empirical expression which describes the variation in leaf photosynthetic capacity (Pm) as a function of the light distribution in the canopy. The light distribution itself was calculated with a simple model which assumes only three leaf angle classes (0–30°, 30–60° and 60–90°). The leaf area index was determined indirectly from measurements of direct radiation below the canopy. The result was a model for canopy photosynthesis that requires only a few parameters. These parameters are the leaf photosynthetic capacity at the top of the canopy, the relative frequency of leaves in each of the three leaf angle classes, and the fraction of direct radiation below the canopy. Each of these parameters can be determined by means of simple non-destructive measurements. The model was applied to dense stands of two monocotyledonous species: rice (Oryza sativa L.) and pearl millet (Pennisetum americanum (L.) K. Schum.). The rates of canopy photosynthesis thus calculated were compared to those obtained with a more elaborate reference model. The differences between the values obtained with the two models were small. The present photosynthesis model can, therefore, be considered to be a suitable alternative for the more elaborate model. It was further discussed that, since the model is based on purely non-destructive measurements, it will be particularly useful in cases where it is required to estimate canopy photosynthesis at regular intervals over a length of time or in stands of vegetation that cover large areas of land.  相似文献   

17.
Changes in canopy structure parameters (leaflet orientation, leaflet inclination and leaf area index) were measured in crops of beans (Phaseolus vulgaris L.) in the field as the canopy developed between July and October. These changes were compared with the corresponding changes in seasonal light transmission. The beans showed clear heliotropic behaviour, with preferential orientation of leaflets towards the sun’s beam, especially on sunny days. Nevertheless a significant proportion of the leaves pointed in other directions, with as much as 20% oriented towards the north. The highest proportion of leaf inclinations was in the range 30–40° on cloudy days and between 40° and 50° on sunny days. Two methods were compared for assessing changes in light transmission: (a) the use of a Sunfleck Ceptometer and (b) the use of continuous records obtained with sensors installed in the canopy. Over the growth period studied, the total of the leaf plus stem area indices (L S) increased from 0.26 to 5.2 with the transmission coefficient (τ) for photosynthetic photon flux density (PPFD), obtained using the Ceptometer, correspondingly decreasing from 0.72 to 0.05, and the canopy extinction coefficient decreasing from 1.4 to 0.62. The continuous records of light transmission gave generally similar estimates of τ. Some contrasting leaf angle distribution functions were compared for estimation of L S from the light measurements. The best leaf angle function to predict L S from the observed light transmission was a conical function corrected by the degree of heliotropism. Received: 27 January 1999 / Accepted: 11 June 1999  相似文献   

18.
Poplars (Populus spp.) have a particular petiole morphology that enhances leaf flutter even in light winds. Previous studies have shown that this trait enhances whole canopy carbon gain through changes in the temporal dynamics and spatial distribution of light in the lower canopy. However, less is known about the effects of flutter for leaves at the top of the canopy ("sun leaves"). A computer simulation model was developed that uses latitude, time of day, day of year, azimuth and a slope component, which was varied at a 3 Hz frequency over an arc of rotation to create the flutter motion, and generate data on light interception for both surfaces of a fixed or fluttering leaf. The light files generated (10 Hz) were input into a dynamic model of photosynthesis to estimate the carbon gain for both fluttering and fixed leaves. As compared to leaves fixed at various angles and azimuths, fluttering leaves had a more uniform light interception. Depending on their angle and azimuth, fixed leaves may not always be intercepting high photon flux density (PFD) even when exposed to full sun. Leaf flutter continuously randomizes leaf angles creating uniform PFD inputs for photosynthetic reactions regardless of variation in leaf orientation and solar position. These effects on light interception could have positive impacts on carbon gain for leaves at the top of the canopy.  相似文献   

19.
Evolutionarily stable leaf area production in plant populations   总被引:4,自引:0,他引:4  
Using an analytical model, it was shown that for a given amount of nitrogen in the canopy of a stand (N(T)), there exists an evolutionarily stable leaf area index (ES-LAI), and therefore an evolutionarily stable average leaf nitrogen content (n(ES)(av);n(ES)(av) =N(T)/ES-LAI), at which no individual plant in the stand can increase its photosynthesis by changing its leaf area. It was also shown that this ES-LAI is always greater than the optimal LAI that maximizes photosynthesis per unit N(T) of the stand. This illustrates that the canopy structure that maximizes photosynthesis of a population is not the same as the canopy structure that maximizes photosynthesis of individuals within a population. It was further derived that the ES-LAI at given N(T) increases with the ratio between the light-saturated photosynthesis and the N content per unit leaf area (leaf-PPNUE) and that it decreases with the canopy extinction coefficient for light (K(L)), the light availability and the apparent quantum yield (phi). These hypotheses were tested by comparing calculated ES-LAI and n(ES)(av) values to actual LAIs and leaf N contents measured for stands of a large variety of herbaceous plants. There was a close correspondence between the calculated and measured values. As predicted by the model, plants with high leaf-PPNUEs produced more leaf area per unit nitrogen than those with low leaf-PPNUEs while plants with horizontal leaves, forming stands with higher K(L) values, produced less leaf area than those with more vertically inclined leaves. These results suggest that maximization of individual plant photosynthesis per unit of nitrogen plays an important role in determining leaf area production of plants and the resulting canopy structure of stands of vegetation. They further suggest this optimization to be a mechanism by which leaf traits such as leaf-PPNUE and leaf inclination angle are causally related to structural characteristics of the population, i.e. the leaf area index of the stand.  相似文献   

20.
Impact of light limitation on seagrasses   总被引:2,自引:0,他引:2  
Seagrass distribution is controlled by light availability, especially at the deepest edge of the meadow. Light attenuation due to both natural and anthropogenically-driven processes leads to reduced photosynthesis. Adaptation allows seagrasses to exist under these sub-optimal conditions. Understanding the minimum quantum requirements for growth (MQR) is revealed when light conditions are insufficient to maintain a positive carbon balance, leading to a decline in seagrass growth and distribution. Respiratory demands of photosynthetic and non-photosynthetic tissues strongly influence the carbon balance, as do resource allocations between above- and below-ground biomass. Seagrass light acclimation occurs on varying temporal scales, as well as across spatial scales, from the position along a single leaf blade to within the canopy and finally across the meadow. Leaf absorptance is regulated by factors such as pigment content, morphology and physical properties. Chlorophyll content and morphological characteristics of leaves such as leaf thickness change at the deepest edge. We present a series of conceptual models describing the factors driving the light climate and seagrass responses under current and future conditions, with special attention on the deepest edge of the meadow.  相似文献   

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