Vegetation clumping modulates global photosynthesis through adjusting canopy light environment

The spatial dispersion of photoelements within a vegetation canopy, quantified by the clumping index (CI), directly regulates the within-canopy light environment and photosynthesis rate, but is not commonly implemented in terrestrial biosphere models to estimate the ecosystem carbon cycle. A few global CI products have been developed recently with remote sensing measurements, making it possible to examine the global impacts of CI. This study deployed CI in the radiative transfer scheme of the Community Land Model version 5 (CLM5) and used the revised CLM5 to quantitatively evaluate the extent to which CI can affect canopy absorbed radiation and gross primary production (GPP), and for the first time, considering the uncertainty and seasonal variation of CI with multiple remote sensing products. Compared to the results without considering the CI impact, the revised CLM5 estimated that sunlit canopy absorbed up to 9%–15% and 23%–34% less direct and diffuse radiation, respectively, while shaded canopy absorbed 3%–18% more diffuse radiation across different biome types. The CI impacts on canopy light conditions included changes in canopy light absorption, and sunlit–shaded leaf area fraction related to nitrogen distribution and thus the maximum rate of Rubisco carboxylase activity (V_cmax), which together decreased photosynthesis in sunlit canopy by 5.9–7.2 PgC year⁻¹ while enhanced photosynthesis by 6.9–8.2 PgC year⁻¹ in shaded canopy. With higher light use efficiency of shaded leaves, shaded canopy increased photosynthesis compensated and exceeded the lost photosynthesis in sunlit canopy, resulting in 1.0 ± 0.12 PgC year⁻¹ net increase in GPP. The uncertainty of GPP due to the different input CI datasets was much larger than that caused by CI seasonal variations, and was up to 50% of the magnitude of GPP interannual variations in the tropical regions. This study highlights the necessity of considering the impacts of CI and its uncertainty in terrestrial biosphere models.     

The impact of diffuse sunlight on canopy light-use efficiency, gross photosynthetic product and net ecosystem exchange in three forest biomes

For three forest canopies (a sparse, boreal needleleaf; a temperate broadleaf; and a dense, tropical, broadleaf stand) light‐use efficiency (LUE) is found to be 6–33% higher when sky radiance is dominated by diffuse rather than direct sunlight. This enhancement is much less than that reported previously for both crops (110%; Choudbury, 2001 ) and moderately dense temperate woodland (50–180%). We use the land‐surface scheme JULES to interpret the observed canopy response. Once sunflecks and leaf orientation are incorporated explicitly into the scheme, our simulations reproduce convincingly the overall level of canopy gross photosynthetic product (GPP), its enhancement with respect to diffuse sunlight and the mean 15% reduction in productivity observed during the afternoon due to stomatal closure. The LUE enhancement under diffuse sunlight can be explained by sharing of the canopy radiation‐load, which is reduced under direct sky radiance. Once sunflecks are accounted for the advantage of implementing more sophisticated calculations of stomatal conductance (e.g. Ball–Berry and SPA submodels) is less obvious even for afternoon assimilation. Empirical relations are developed between observed carbon flux and the environmental variables total downwelling shortwave radiation (SW), canopy temperature (T) and the fraction of diffuse sky radiance (fDIF). These relations allow us to gauge the impact of increased/reduced insolation on GPP and net ecosystem exchange (NEE). Overall the three stands appear to be fairly stable within global trends and typical interannual variability (SW changing by <15%). Greatest sensitivity is exhibited by the boreal site, Zotino, where NEE falls by 9±4% for a 15% reduction in SW.     

美国黑核桃实生苗生态生理过程对环境因素响应的数值模拟(Ⅲ):植株冠层光合作用数理模型

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．     

Effects of seasonal and interannual variations in leaf photosynthesis and canopy leaf area index on gross primary production of a cool-temperate deciduous broadleaf forest in Takayama, Japan

Revealing the seasonal and interannual variations in forest canopy photosynthesis is a critical issue in understanding the ecological mechanisms underlying the dynamics of carbon dioxide exchange between the atmosphere and deciduous forests. This study examined the effects of temporal variations of canopy leaf area index (LAI) and leaf photosynthetic capacity [the maximum velocity of carboxylation (V _cmax)] on gross primary production (GPP) of a cool-temperate deciduous broadleaf forest for 5 years in Takayama AsiaFlux site, central Japan. We made two estimations to examine the effects of canopy properties on GPP; one is to incorporate the in situ observation of V _cmax and LAI throughout the growing season, and another considers seasonality of LAI but constantly high V _cmax. The simulations indicated that variation in V _cmax and LAI, especially in the leaf expansion period, had remarkable effects on GPP, and if V _cmax was assumed constant GPP will be overestimated by 15%. Monthly examination of air temperature, radiation, LAI and GPP suggested that spring temperature could affect canopy phenology, and also that GPP in summer was determined mainly by incoming radiation. However, the consequences among these factors responsible for interannual changes of GPP are not straightforward since leaf expansion and senescence patterns and summer meteorological conditions influence GPP independently. This simulation based on in situ ecophysiological research suggests the importance of intensive consideration and understanding of the phenology of leaf photosynthetic capacity and LAI to analyze and predict carbon fixation in forest ecosystems.     

Ecophysiological controls over the net ecosystem exchange of mountain spruce stand. Comparison of the response in direct vs. diffuse solar radiation

Cloud cover increases the proportion of diffuse radiation reaching the Earth's surface and affects many microclimatic factors such as temperature, vapour pressure deficit and precipitation. We compared the relative efficiencies of canopy photosynthesis to diffuse and direct photosynthetic photon flux density (PPFD) for a Norway spruce forest (25‐year‐old, leaf area index 11 m² m⁻²) during two successive 7‐day periods in August. The comparison was based on the response of net ecosystem exchange (NEE) of CO₂ to PPFD. NEE and stomatal conductance at the canopy level (G_canopy) was estimated from half‐hourly eddy‐covariance measurements of CO₂ and H₂O fluxes. In addition, daily courses of CO₂ assimilation rate (A_N) and stomatal conductance (G_s) at shoot level were measured using a gas‐exchange technique applied to branches of trees. The extent of spectral changes in incident solar radiation was assessed using a spectroradiometer. We found significantly higher NEE (up to 150%) during the cloudy periods compared with the sunny periods at corresponding PPFDs. Prevailing diffuse radiation under the cloudy days resulted in a significantly lower compensation irradiance (by ca. 50% and 70%), while apparent quantum yield was slightly higher (by ca. 7%) at canopy level and significantly higher (by ca. 530%) in sun‐acclimated shoots. The main reasons for these differences appear to be (1) more favourable microclimatic conditions during cloudy periods, (2) stimulation of photochemical reactions and stomatal opening via an increase of blue/red light ratio, and (3) increased penetration of light into the canopy and thus a more equitable distribution of light between leaves. Our analyses identified the most important reason of enhanced NEE under cloudy sky conditions to be the effective penetration of diffuse radiation to lower depths of the canopy. This subsequently led to the significantly higher solar equivalent leaf area compared with the direct radiation. Most of the leaves in such dense canopy are in deep shade, with marginal or negative carbon balances during sunny days. These findings show that the energy of diffuse, compared with direct, solar radiation is used more efficiently in assimilation processes at both leaf and canopy levels.     

A new paradigm in leaf-level photosynthesis: direct and diffuse lights are not equal

Global-change scenarios suggest a trend of increasing diffuse light due to expected increases in cloud cover. Canopy-level measurements of plant-community photosynthesis under diffuse light show increased productivity attributed to more uniform distribution of light within the forest canopy, yet the effect of the directional quality of light at the leaf level is unknown. Here we show that leaf-level photosynthesis in sun leaves of both C(3) and C(4) plants can be 10-15% higher under direct light compared to equivalent absorbed irradiances of diffuse light. High-light-grown leaves showed significant photosynthetic enhancement in direct light, while shade-adapted leaves showed no preference for direct or diffuse light at any irradiance. High-light-grown leaves with multiple palisade layers may be adapted to better utilize direct than diffuse light, while shade leaf structure does not appear to discriminate light based on its directionality. Based upon our measurements, it appears that leaf-level and canopy-level photosynthetic processes react differently to the directionality of light, and previously observed increases in canopy-level photosynthesis occur even though leaf-level photosynthesis decreases under diffuse light.     

Spectral composition of photosynthetically active radiation penetrating into a Norway spruce canopy: the opposite dynamics of the blue/red spectral ratio during clear and overcast days

The spectral composition of photosynthetically active radiation (PAR) during clear and overcast days was studied above the canopy (U) and at two layers of a dense Norway spruce stand [Picea abies (L.) Karst.] characterized with an average LAI = 7.3 ± 0.8 (middle layer: M) and 12.3 ± 0.7 (lower layer: L). Whereas the spectral composition of PAR incoming on the canopy surface during cloudy days (characterized by diffuse index DI > 0.7) was almost independent of the solar elevation angle, the proportion of the blue-green spectral region of PAR was significantly reduced at low elevation angles during days with prevailing direct radiation (DI < 0.3). The PAR spectrum at both M and L levels was only slightly enriched in the green spectral region (more pronounced for DI < 0.3). The penetration of diffuse radiation into the canopy resulted in a slight (approx. 5%) reduction of the blue region proportion that remained stable during the day. On the contrary, under clear sky conditions the penetration of blue and red radiation was dependent on the solar elevation in an opposite manner in comparison with the spectral composition of PAR incident on canopy, giving almost twofold proportion of the blue part of the spectrum at a low elevation angle at M layer. We suggest that the blue enhancement of the spectrum within the Norway spruce canopy during clear days is due to a specific spatial arrangement of the assimilatory apparatus of a coniferous stand. Further, the possible consequences of the observed dynamics of the PAR spectrum inside the canopy during clear days on the efficiency of PAR absorption of the needles located within the canopy are discussed.     

Chlorophyll fluorescence tracks seasonal variations of photosynthesis from leaf to canopy in a temperate forest

Accurate estimation of terrestrial gross primary productivity (GPP) remains a challenge despite its importance in the global carbon cycle. Chlorophyll fluorescence (ChlF) has been recently adopted to understand photosynthesis and its response to the environment, particularly with remote sensing data. However, it remains unclear how ChlF and photosynthesis are linked at different spatial scales across the growing season. We examined seasonal relationships between ChlF and photosynthesis at the leaf, canopy, and ecosystem scales and explored how leaf‐level ChlF was linked with canopy‐scale solar‐induced chlorophyll fluorescence (SIF) in a temperate deciduous forest at Harvard Forest, Massachusetts, USA. Our results show that ChlF captured the seasonal variations of photosynthesis with significant linear relationships between ChlF and photosynthesis across the growing season over different spatial scales (R² = 0.73, 0.77, and 0.86 at leaf, canopy, and satellite scales, respectively; P < 0.0001). We developed a model to estimate GPP from the tower‐based measurement of SIF and leaf‐level ChlF parameters. The estimation of GPP from this model agreed well with flux tower observations of GPP (R² = 0.68; P < 0.0001), demonstrating the potential of SIF for modeling GPP. At the leaf scale, we found that leaf F_q’/F_m’, the fraction of absorbed photons that are used for photochemistry for a light‐adapted measurement from a pulse amplitude modulation fluorometer, was the best leaf fluorescence parameter to correlate with canopy SIF yield (SIF/APAR, R² = 0.79; P < 0.0001). We also found that canopy SIF and SIF‐derived GPP (GPP_SIF) were strongly correlated to leaf‐level biochemistry and canopy structure, including chlorophyll content (R² = 0.65 for canopy GPP_SIF and chlorophyll content; P < 0.0001), leaf area index (LAI) (R² = 0.35 for canopy GPP_SIF and LAI; P < 0.0001), and normalized difference vegetation index (NDVI) (R² = 0.36 for canopy GPP_SIF and NDVI; P < 0.0001). Our results suggest that ChlF can be a powerful tool to track photosynthetic rates at leaf, canopy, and ecosystem scales.     

Early-season effects of supplemented solar UV-B radiation on seedling emergence, canopy structure, simulated stand photosynthesis and competition for light

Mixtures and monocultures of wheat (Triticum aestivum) and wild oat (Avena fatua), a common weedy competitor of wheat, were exposed to enhanced solar UV-B radiation simulating a 20% reduction in stratospheric ozone to assess the timing and seasonal development of the UV-B effects on light competition in these species. Results from two years of field study revealed that UV-B enhancement had no detectable effect on the magnitude or timing of seedling emergence in either species. End-of-season measurements showed significant UV-B inhibition of leaf insertion height in wild oat in mixture and monoculture in the second year (irrigated year) but not in the first year (drought year). Leaf insertion height of wheat was not affected by UV-B in either year. The UV-B treatment had no detectable effect on monoculture or total (combined species) mixture LAI but did significantly increase (5–7%) the fractional contribution of wheat to the mixture LAI after four weeks of growth in both years. In addition, the UV-B treatment had subtle effects on LAI height profiles with early season mixtures showing significant reductions in wild oat LAI in lower canopy layers in both years while midseason Year 2 mixtures showed significant reductions in wild oat LAI in upper canopy layers. The changes in canopy structure were found to significantly increase (6–7%) the proportional simulated clear sky canopy photosynthesis and light interception of wheat in mixture. These findings, and others, indicate that the effects of UV-B enhancement on competition are realized very early in canopy development and provide additional support for the hypothesis that UV-B enhancement may shift the balance of competition between these species indirectly by altering competitive interactions for light.     

Environmental controls on the light use efficiency of terrestrial gross primary production

Gross primary production (GPP) by terrestrial ecosystems is a key quantity in the global carbon cycle. The instantaneous controls of leaf-level photosynthesis are well established, but there is still no consensus on the mechanisms by which canopy-level GPP depends on spatial and temporal variation in the environment. The standard model of photosynthesis provides a robust mechanistic representation for C₃ species; however, additional assumptions are required to “scale up” from leaf to canopy. As a consequence, competing models make inconsistent predictions about how GPP will respond to continuing environmental change. This problem is addressed here by means of an empirical analysis of the light use efficiency (LUE) of GPP inferred from eddy covariance carbon dioxide flux measurements, in situ measurements of photosynthetically active radiation (PAR), and remotely sensed estimates of the fraction of PAR (fAPAR) absorbed by the vegetation canopy. Focusing on LUE allows potential drivers of GPP to be separated from its overriding dependence on light. GPP data from over 100 sites, collated over 20 years and located in a range of biomes and climate zones, were extracted from the FLUXNET2015 database and combined with remotely sensed fAPAR data to estimate daily LUE. Daytime air temperature, vapor pressure deficit, diffuse fraction of solar radiation, and soil moisture were shown to be salient predictors of LUE in a generalized linear mixed-effects model. The same model design was fitted to site-based LUE estimates generated by 16 terrestrial ecosystem models. The published models showed wide variation in the shape, the strength, and even the sign of the environmental effects on modeled LUE. These findings highlight important model deficiencies and suggest a need to progress beyond simple “goodness of fit” comparisons of inferred and predicted carbon fluxes toward an approach focused on the functional responses of the underlying dependencies.     

Development of the Monsi-Saeki theory on canopy structure and function

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.     

Exploring the spatial distribution of light interception and photosynthesis of canopies by means of a functional-structural plant model

Background and Aims

At present most process-based models and the majority of three-dimensional models include simplifications of plant architecture that can compromise the accuracy of light interception simulations and, accordingly, canopy photosynthesis. The aim of this paper is to analyse canopy heterogeneity of an explicitly described tomato canopy in relation to temporal dynamics of horizontal and vertical light distribution and photosynthesis under direct- and diffuse-light conditions.

Methods

Detailed measurements of canopy architecture, light interception and leaf photosynthesis were carried out on a tomato crop. These data were used for the development and calibration of a functional–structural tomato model. The model consisted of an architectural static virtual plant coupled with a nested radiosity model for light calculations and a leaf photosynthesis module. Different scenarios of horizontal and vertical distribution of light interception, incident light and photosynthesis were investigated under diffuse and direct light conditions.

Key Results

Simulated light interception showed a good correspondence to the measured values. Explicitly described leaf angles resulted in higher light interception in the middle of the plant canopy compared with fixed and ellipsoidal leaf-angle distribution models, although the total light interception remained the same. The fraction of light intercepted at a north–south orientation of rows differed from east–west orientation by 10 % on winter and 23 % on summer days. The horizontal distribution of photosynthesis differed significantly between the top, middle and lower canopy layer. Taking into account the vertical variation of leaf photosynthetic parameters in the canopy, led to approx. 8 % increase on simulated canopy photosynthesis.

Conclusions

Leaf angles of heterogeneous canopies should be explicitly described as they have a big impact both on light distribution and photosynthesis. Especially, the vertical variation of photosynthesis in canopy is such that the experimental approach of photosynthesis measurements for model parameterization should be revised.     

Developing a diagnostic model for estimating terrestrial vegetation gross primary productivity using the photosynthetic quantum yield and Earth Observation data

This article develops a new carbon exchange diagnostic model [i.e. Southampton CARbon Flux (SCARF) model] for estimating daily gross primary productivity (GPP). The model exploits the maximum quantum yields of two key photosynthetic pathways (i.e. C₃ and C₄) to estimate the conversion of absorbed photosynthetically active radiation into GPP. Furthermore, this is the first model to use only the fraction of photosynthetically active radiation absorbed by photosynthetic elements of the canopy (i.e. FAPAR_ps) rather than total canopy, to predict GPP. The GPP predicted by the SCARF model was comparable to in situ GPP measurements (R² > 0.7) in most of the evaluated biomes. Overall, the SCARF model predicted high GPP in regions dominated by forests and croplands, and low GPP in shrublands and dry‐grasslands across USA and Europe. The spatial distribution of GPP from the SCARF model over Europe and conterminous USA was comparable to those from the MOD17 GPP product except in regions dominated by croplands. The SCARF model GPP predictions were positively correlated (R² > 0.5) to climatic and biophysical input variables indicating its sensitivity to factors controlling vegetation productivity. The new model has three advantages, first, it prescribes only two quantum yield terms rather than species specific light use efficiency terms; second, it uses only the fraction of PAR absorbed by photosynthetic elements of the canopy (FAPAR_ps) hence capturing the actual PAR used in photosynthesis; and third, it does not need a detailed land cover map that is a major source of uncertainty in most remote sensing based GPP models. The Sentinel satellites planned for launch in 2014 by the European Space Agency have adequate spectral channels to derive FAPAR_ps at relatively high spatial resolution (20 m). This provides a unique opportunity to produce global GPP operationally using the Southampton CARbon Flux (SCARF) model at high spatial resolution.     

Solar ultraviolet-B and photosynthetically active irradiance in the urban sub-canopy: A survey of influences

Stratospheric ozone loss in mid-latitudes is expected to increase the ultraviolet-B (UVB) radiation at the earth's surface. Impacts of this expected increase will depend on many factors, including the distribution of light in other wavelengths. Measurements of the photosynthetically active radiation (PAR) and UVB irradiance were made under clear skies at an open field and under the canopy of scattered trees in a suburban area in W. Lafayette, Indiana, USA (latitude 40.5°). Results showed that when there was significant sky view, the UVB penetration into sub-canopy spaces differs greatly from that of PAR. The UVBT _canopy (transmittance; irradiance below canopy/irradiance in open) was inversely related to sky view. The UVB irradiance did not vary as greatly between shaded and sunlit areas as did PAR. Analysis of measurements made near a brick wall indicated that the leaf area of a canopy and the brick wall primarily acted to block fractions of the sky radiance and contributed little scattered UVB to the horizontal plant. A model was developed to predict the UVB and PART _canopy based on diffuse fraction, sky view, and porosity of the crown(s) through which the beam is penetrating. The model accounted for the UVB and PART _canopy to within 0.13 and 0.05 root mean squared error (RMSE), respectively. Analysis of the errors due to model assumptions indicated that care must be taken in describing the sky radiance distribution, the porosity of trees, the penetration of diffuse radiation through porous trees, and the location of sky-obstructing trees and buildings.     

小麦LAI-2000观测值对辐亮度变化的响应

利用LAI-2000植物冠层分析仪和ASD光谱仪,通过固定点拔节期冬小麦叶面积指数(LAI)观测实验和同步光谱辐亮度实验,测量了晴天条件下,固定点冬小麦从中午至傍晚24个不同时刻的LAI值及对应的辐亮度。继之分析了此段时间内测得的定点冬小麦LAI值分别与对应时刻可见光和近红外谱段的天空光辐亮度、总辐亮度和太阳直射辐亮度值之间的相关性。结果表明,无论波长小于490nm的谱段,还是波长大于490 nm的谱段,LAI与天空光辐亮度、总辐亮度和太阳直射辐亮度都呈负相关,相关系数(R2)高达0.8左右;尤其LAI与天空光辐亮度的负相关性最高,这种相关性随着波长的增大而减小。LAI与各谱段天空光辐亮度的相关性特征可为LAI-2000晴天观测的LAI值归一化修正处理提供一种新思路和技术途径,以消除太阳直射光的影响,从而解决LAI-2000只能在晴天观测的局限性,拓展LAI-2000在晴天观测条件下的适用性。     

Canopy structure,light microclimate and leaf gas exchange of Quercus coccifera L. in a Portuguese macchia: measurements in different canopy layers and simulations with a canopy model

Summary The structural characteristics of a diverse array of Quercus coccifera canopies were assessed and related to measured and computed light attenuation, proportion of sunlit foliage, foliage temperatures, and photosynthesis and diffusive conductance behavior in different canopy layers. A canopy model incorporating all components of shortwave and longwave radiation, and the energy balance, conductance, and CO₂ and H₂O exchanges of all leaf layers was developed and compared with measurements of microclimate and gas exchange in canopies in four seasons of the year. In the denser canopies with a leaf area index (LAI) greater than 5, there is little sunlit foliage and the diffuse radiation (400–700 nm) is attenuated to 5% or less of the global radiation (400–700 nm) incident on the top of the canopy. Foliage of this species is nonrandomly distributed with respect to azimuth angle, and within each canopy layer, foliage azimuth and inclination angles are correlated. A detailed version of the model which computed radiation interception and photosynthetic light harvesting according to these nonrandom distributions indicated little difference in whole-canopy gas exchange from calculations of the normal model, which assumes random azimuth orientation. The contributions of different leaf layers to canopy gas exchange are not only a function of the canopy microclimate, but also the degree to which leaves in the lower layers of the canopy exhibit more shade-leaf characteristics, such as low photosynthetic and respiratory capacity and maximal conductance. On cloudless days, the majority of the foliage in a canopy of 5.4 LAI is shaded —70%–90% depending on the time of year. Yet, the shaded foliage under these conditions is calculated to contribute only about one-third of the canopy carbon gain. This contribution is about the same as that of the upper 13% of the canopy foliage. Computed annual whole-canopy carbon gain and water use are, respectively, 60% and 100% greater for a canopy of 5 LAI than for one of 2 LAI. Canopy water-use efficiency is correspondingly less for the canopy of 5 LAI than for that of 2 LAI, but most of this difference is apparent during the cool months of the year, when moisture is more abundant.     

Modelling Plant Responses to Elevated CO2: How Important is Leaf Area Index?

BACKGROUND AND AIMS: The problem of increasing CO(2) concentration [CO(2)] and associated climate change has generated much interest in modelling effects of [CO(2)] on plants. While variation in growth and productivity is closely related to the amount of intercepted radiation, largely determined by leaf area index (LAI), effects of elevated [CO(2)] on growth are primarily via stimulation of leaf photosynthesis. Variability in LAI depends on climatic and growing conditions including [CO(2)] concentration and can be high, as is known for agricultural crops which are specifically emphasized in this report. However, modelling photosynthesis has received much attention and photosynthesis is often represented inadequately detailed in plant productivity models. Less emphasis has been placed on the modelling of leaf area dynamics, and relationships between plant growth, elevated [CO(2)] and LAI are not well understood. This Botanical Briefing aims at clarifying the relative importance of LAI for canopy assimilation and growth in biomass under conditions of rising [CO(2)] and discusses related implications for process-based modelling. MODEL: A simulation exercise performed for a wheat crop demonstrates recent experimental findings about canopy assimilation as affected by LAI and elevation of [CO(2)]. While canopy assimilation largely increases with LAI below canopy light saturation, effects on canopy assimilation of [CO(2)] elevation are less pronounced and tend to decline as LAI increases. Results from selected model-testing studies indicate that simulation of LAI is often critical and forms an important source of uncertainty in plant productivity models, particularly under conditions of limited resource supply. CONCLUSIONS: Progress in estimating plant growth and productivity under rising [CO(2)] is unlikely to be achieved without improving the modelling of LAI. This will depend on better understanding of the processes of substrate allocation, leaf area development and senescence, and the role of LAI in controlling plant adaptation to environmental changes.     

Estimation of deciduous forest leaf area index using direct and indirect methods

This study evaluated one semi-direct and three indirect methods for estimating leaf area index (LAI) by comparing these estimates with direct estimates derived from litter collection. The semi-direct method uses a thin metallic needle to count a number of contacts across fresh litter layers. One indirect method is based on the penetration of diffuse global radiation measured over the course of a day. The second indirect method uses the LAI-2000 plant canopy analyser (PCA) which measures diffuse light penetration from five different sky sectors simultaneously. The third indirect method uses the Demon portable light sensor to measure the penetration of direct beam sunlight at different zenith angles over the course of half a day. The Poisson model of gap frequency was applied to estimate plant area index (PAI) from observed transmittances using the second and third methods. Litter collection from 11 temperate decidous forests gave values of LAI ranging from 1.7 to 7.5. Estimates based on the needle method showed a significant linear relationship with LAI values obtained from litter collections but were systematically lower (by 6–37%). PAI estimates using all three indirect techniques (fixed light sensor system, LAI-2000 and Demon) showed a strong linear relationship with LAI derived from litter collection. Differences, averaged over all forest stands, between PAI estimates from each of the three indirect methods and LAI from litter collections were below 2%. If we consider that LAI=PAI–WAI (wood area index) then, all three indirect methods underestimated LAI by an additional factor close to the value of WAI. One reason could be a local clumping of architectural canopy components: in particular, the spatial dispositions of branchlets and leaves are not independent, leading to a non-random relationship between the distributions of these two canopy components.     

Daylight spectra (400–740 nm) beneath sunny,blue skies in Tasmania,and the effect of a forest canopy

Abstract The photosynthetically active radiation (PAR) incident on a horizontal surface at an open mountain site is positively correlated with solar altitude for sunny, blue sky conditions. The proportion of red light in PAR decreases with increasing solar altitude, while that of blue increases. These results are consistent with the wavelength dependency of Rayleigh and Mie scattering. The ratio of near infrared radiation to PAR decreases with increasing solar altitude towards solar noon and with decreasing solar altitude towards sunset. Thus surface reflection seems to be an important part of the light climate. The relative transmission of daylight through a forest canopy to a horizontal surface is not correlated with solar altitude for sunny, blue sky conditions at a mountain site. The amount of diffuse daylight is negatively correlated with per cent canopy interception, and the amount of direct sunlight is negatively correlated with per cent solar track interception. Daylength is negatively correlated with both canopy and solar track interceptions. The proportion of red light in PAR increases with increasing solar altitude, while that of blue decreases. These results are opposite those for the open site and are due to the spatial patterns of canopy obstruction of the sky vault, and of the spectral quality of daylight across the sky. The ratio of near infrared radiation to PAR in shadelight increases with increasing canopy interception due to the selective scattering properties of the canopy. The ratio for shadelight is positively correlated with the ratio for sunflecks.     

Improving the light use efficiency model for simulating terrestrial vegetation gross primary production by the inclusion of diffuse radiation across ecosystems in China

Qualification of gross primary production (GPP) of terrestrial ecosystem over large areas is important in understanding the response of terrestrial ecosystem to global climate change. While light use efficiency (LUE) models were widely used in regional carbon budget estimates, few studies consider the effect of diffuse radiation on LUE caused by clouds using a big leaf model. Here we developed a cloudiness index light use efficiency (CI-LUE) model based on the MOD17 model algorithm to estimate the terrestrial ecosystem GPP, in which the base light use efficiency encompassed the cloudiness index, maximum LUE and clear sky LUE. GPP measured at seven sites from 2003 to 2007 in China were used to calibrate and validate the CI-LUE model. The results showed that at forest sites and cropland site the CI-LUE model outperformed the Vegetation Photosynthesis Model (VPM), Terrestrial Ecosystem Carbon flux model (TEC), MOD17 model algorithm driven by in situ meteorological measurements and MODIS GPP products, especially the R² of simulated GPP against flux measurements at Dinghushan forest site increased from 0.17 (MODIS GPP products) to 0.61 (CI-LUE). Instead, VPM model had the best agreement with GPP measurements followed by CI-LUE model and lastly TEC model at two grassland sites. Meanwhile, GPP calculated by CI-LUE model has less underestimation under cloudy skies in comparison with MOD17 model. This study demonstrated the potential of the CI-LUE model in improving GPP simulations resulting from the inclusion of diffuse radiation in regulating the base light use efficiency and maximum light use efficiency.