落叶阔叶林冠层光合有效辐射分量的遥感模拟与分析

冠层绿色叶片(光合组分)的光合有效辐射分量(绿色FPAR)真实地反映了植被与外界进行物质和能量交换的能力,获取冠层光合组分吸收的太阳光合有效辐射,对生态系统生产力的遥感估算精度的提高具有重要的意义。研究以落叶阔叶林为例,基于SAIL模型模拟森林冠层光合组分和非光合组分吸收的光合有效辐射,研究冠层FPAR变化规律以及与植被指数的相关关系。结果表明,冠层结构的改变会影响冠层对PAR的吸收能力,冠层绿色FPAR的大小与植被面积指数及光合组分面积比相关;在高覆盖度植被区,冠层绿色FPAR占冠层总FPAR的80%以上,非光合组分的贡献较小,但在低植被覆盖区,当光合组分和非光合组分面积相同时,绿色FPAR不及冠层总FPAR的50%;相比于NDVI,北方落叶阔叶林冠层EVI与绿色FPAR存在更为显著的线性相关关系(R~20.99)。     

Gross primary productivity and transpiration flux of the Australian vegetation from 1788 to 1988 AD: effects of CO2 and land use change

We present a novel approach to estimating the transpiration flux and gross primary productivity (GPP) from Normalized Difference Vegetation Index, leaf functional types, and readily available climate data. We use this approach to explore the impact of variations in the concentration of carbon dioxide in the atmosphere ([CO₂]) and consequent predicted changes in vegetation cover, on the transpiration flux and GPP. There was a near 1 : 1 relationship between GPP estimated with this transpiration flux approach and that estimated using a radiation‐use efficiency (RUE) approach. Model estimates are presented for the Australian continent under three vegetation–[CO₂] scenarios: the present vegetation and hypothetical ‘natural’ vegetation cover with atmospheric CO₂ concentration ([CO₂]) of 350 μmol mol^?1 (pveg₃₅₀ and nveg₃₅₀), and for the ‘natural’ vegetation with [CO₂] 280 μmol mol^?1 (nveg₂₈₀). Estimated continental GPP is 6.5, 6.3 and 4.3 Gt C yr^?1 for pveg₃₅₀, nveg₃₅₀ and nveg₂₈₀, respectively_. The corresponding transpiration fluxes are 232, 224 and 190 mm H₂O yr^?1. The contribution of the raingreen and evergreen components of the canopy to these fluxes are also estimated.     

The microclimate under coloured hailnets affects leaf and fruit temperature, leaf anatomy, vegetative and reproductive growth as well as fruit colouration in apple

The purpose of this study was to investigate supposedly positive biological effects of coloured hailnets on microclimate, including photosynthetically active radiation (PAR), UV-B, air, soil, fruit and leaf temperature as well as humidity, which in turn may affect leaf anatomy, tree growth and fruit quality; apple was chosen as a model crop at Klein-Altendorf near Bonn, Germany; adjacent uncovered trees served as control. Red and green hailnets transmitted 3–6% more red or green light, without alteration of the red:far red (R–666 nm:FR–730 nm) ratio (0.99–1.01:1) and hence without affecting the phytochrome system. The microclimate was changed with a reduction by 12–23% in PAR and, to a larger extent, by 20–28% in UV, viz. shading. Light measurements at a 45° angle, to mimic the fruit or leaf position, showed that PAR was 90–210 µmol m⁻² s⁻¹ larger outside on a sunny summer day than under the white or red-white and 150–340 µmol m⁻² s⁻¹ larger than under red-black and green-black hailnets. Air temperature and relative humidity under coloured hailnets decreased by ca. 1.3°C and by ca 2% rh (cloudy) to 5% rh (sunny day), respectively, compared with outside; leaf temperature was decreased by up to 3°C and fruit temperature by up to 6°C. Soil temperatures at 5 cm depth were 0.5–1°C colder under red-black and green-black hailnets, but up to 0.9°C warmer under white and red-white hailnets compared with the uncovered control outside. Alternate bearing had a larger impact on vegetative growth in the affected year than the coloured hailnets; annual trunk diameter increments in cv. ‘Fuji’, i.e. the variety susceptible to alternate bearing, showed a larger variation than cv. ‘Pinova’ without alternate bearing. Reproductive growth, viz. return bloom and leaf anatomy were impaired by the coloured hailnets. Apple trees under dark hailnets developed thinner leaves with a thinner epidermis and fewer layers of palisade cells. These leaves were 3.5°C (dark hailnets) and 2.5°C (white hailnets) cooler than outside on a sunny day compared with ca. 1.5°C (dark hailnets) and 0.85°C (white hailnets) on a cloudy day. Transpirational cooling of cv. ‘Fuji’ leaves was 0.3–0.6°C outside and 1.4–1.6°C under the green-black hailnet on sunny days compared to <0.1°C on cloudy days. As a practical application, apple fruit colouration was dependent on light (PAR and UV-B) transmission of the respective hailnet colour.     

黄土高原不同植被覆被类型NDVI对气候变化的响应

植被与气候是目前研究生态与环境的重要内容。为探究黄土高原地区植被与气候因子之间的响应机制,利用线性趋势分析、Pearson相关分析、多元线性回归模型以及通径分析的方法,对黄土高原2000—2015年全区和不同植被覆被类型区内NDVI与气候因子的变化趋势以及相互作用关系进行分析。植被覆被分类数据和植被指数数据分别来源于ESA CCI-LC(The European Space Agency Climate Change Initiative Land Cover)以及MODND1T/NDVI(Normalized Difference Vegetation Index)。结果表明:(1) 2000—2015年黄土高原全区植被年NDVI_(max)显著增加的区域占总面积的74.25%,不同植被覆被类型年NDVI_(max)分别为常绿阔叶林常绿针叶林落叶阔叶林落叶针叶林镶嵌草地农田镶嵌林地草地灌木,并且都呈显著增加趋势,其中常绿阔叶林和农田增加幅度最大,为0.012/a。(2)黄土高原全区NDVI与气温、日照、降水和相对湿度等气候因子之间没有显著相关性,但在不同植被覆被类型区,气候因子对NDVI存在显著作用,且不同植被覆被类型差异明显。(3)在全区和不同植被覆被类型区NDVI仅对降水的响应比较一致,气温无论在整个区域尺度还是不同植被覆被类型区对植被的影响均不显著。(4)常绿阔叶林、落叶阔叶林、常绿针叶林及镶嵌林地等以乔木为主的植被覆被类型受年均相对湿度和年总日照时数的显著负效应驱动,草地、镶嵌草地等以草本为主的植被覆被类型则受到年总降水量的显著正效应影响。这说明对植被类型进行区分,更有利于揭示气候对植被的作用机制。     

Number of genes controlling slow rusting resistance to leaf rust in five spring wheat cultivars

The number of genes controlling slow rusting resistance to leaf rust (Puccinia triticina) was estimated in five spring wheat (Triticum aestivum) cultivars using quantitative formulae. Parents and F₆ families were evaluated in replicated field trials under epidemics initiated by artificial inoculation. The F₆ families resulted from a diallel cross involving the fast-rusting cultivar Yecora 70 and five slow-rusting wheat cultivars: Sonoita 81, Tanager ‘S’, Galvez 87, Ures 81, and Moncho ‘S’. The area under the disease progress curve (AUDPC) was used to measure leaf rust severity over time. Results indicate that cultivar Sonoita 81 has three or four genes, Tanager ‘S’ has two or three genes, Galvez 87 has three genes, and both Ures 81 and Moncho ‘S’ have two genes for slow rusting resistance to leaf rust. Based on this result and previously reported moderate to high narrow-sense heritability estimates for slow rusting resistance in these materials, early-generation selection for slow leaf rusting would be effective.     

低温胁迫对水稻幼苗不同叶龄叶片叶绿素荧光特性的影响

以‘蜀恢162’(‘Shuhui 162’)、‘糯89-1’(‘Nuo 89-1’)、‘蜀恢162/糯89-1’(‘Shuhui 162/Nuo 89-1’)、‘奇妙香’(‘Qimiaoxiang’)和早黄矮(‘Zaohuang’ai’)5个水稻(Oryza sativa L.)品种(系)为研究对象,采用叶绿素荧光成像系统研究了低温(4℃)胁迫对水稻3叶期幼苗不同叶龄叶片叶绿素荧光特性的影响。结果表明:经低温胁迫处理后,5个水稻品种(系)幼苗3个叶龄叶片的各叶绿素荧光参数变化有明显差异,其中第一叶的各项参数均降至0。经低温处理后5个水稻品种(系)幼苗3片叶片的PSⅡ最大光化学量子产量(Fv/Fm)均明显小于对照(25℃),其中第一叶的降低幅度最大、第三叶最小。经低温胁迫处理后,5个水稻品种(系)幼苗第三叶的非光化学淬灭系数(qN)均显著大于对照,耐冷性品种‘糯89-1’幼苗第二叶的qN较对照显著增大,而其他水稻品种(系)幼苗第二叶的qN均显著小于对照;‘糯89-1’幼苗第二叶的光化学淬灭系数(qP)较对照略有增大,第三叶的qP显著大于对照;‘早黄矮’幼苗第三叶的qP也大于对照但差异不显著,而其余水稻品种(系)幼苗第二叶和第三叶的qP均显著小于对照。经低温胁迫后5个水稻品种(系)幼苗3片叶片的PSⅡ最大相对电子传递速率(rETRmax)和半饱和光强(Ik)均显著小于对照;除‘糯89-1’幼苗第三叶外,5个水稻品种(系)幼苗3片叶片的快速光响应曲线初始斜率(α)也均显著小于对照,总体上第一叶的rETRmax、Ik和α下降幅度最大、第三叶最小。研究结果揭示:受低温胁迫后,叶片自身生理差异是导致水稻幼苗不同叶龄叶片受伤害程度不同的主要因素。     

Increase in light interception cost and metabolic mass component of leaves are coupled for efficient resource use in the high altitude vegetation

Global syntheses of leaf trait scaling relationships report an increase in light interception costs or ‘diminishing returns’ with increase in leaf area. However, variation in light interception costs across ecological gradients and plant strategies to cope up with these costs are not adequately understood. We analyzed leaf area (A) – leaf dry mass (M), leaf water mass (W) – M and W – A scaling relationships in plants occurring in a high altitude region of western Himalaya across environmental gradients to understand changes in light interception cost and metabolic mass component. M represents light interception cost, whereas, W is considered as a proxy of metabolic mass component for liquid phase being the ultimate source of metabolic activity. Trait values were measured from 9278 leaves belonging to 136 dominant species occurring at different sites, slope aspects, elevations and habitat types. Overall, light interception cost increased with increasing A (scaling exponent (α) < 1 in A–M relationship) and metabolic mass component increased disproportionately high with increasing M and A. We found significant differences in scaling exponents of leaf trait relationship between sites, elevations, slope aspects and habitat types, indicating that increase in light interception cost was more evident at higher elevations, southern slopes and open habitats. Further, with increase in light interception cost, metabolic mass component also increased (α > 1 in W–M and W–A relationships). The changes in scaling exponents of various leaf trait relationships across ecological gradients indicated that vegetation of different regions have differences in light interception cost and metabolic mass component. Moreover, increasing light interception cost (increase in mechanical and hydraulic tissues) with increasing A and increasing metabolic mass (leaf thickness) with increasing A and M are favored in high altitude vegetation. This could be a key strategy of high altitude plants for efficient resource capture and use in harsh environments.     

Global climate models and 'dynamic' vegetation changes

Models of global change must come to incorporate changes in terrestrial vegetation. Here we choose a 1- year meshing (coupling) period to link a global climate model to a well-known biophysical representation of the continental surface by means of eleven vegetation functional types. This coupled model is used to answer two questions: Can a ‘standard’ GCM ‘cope' with sudden switches in continental characteristics?’ and Does the climate ‘care’ about the changing underlying vegetation? We find affirmative answers to both questions. Our results also suggest that those content to generate vegetation post facto from climate output have incomplete results.     

The influence of leaf size and shape on leaf thermal dynamics: does theory hold up under natural conditions?

Laboratory studies on artificial leaves suggest that leaf thermal dynamics are strongly influenced by the two‐dimensional size and shape of leaves and associated boundary layer thickness. Hot environments are therefore said to favour selection for small, narrow or dissected leaves. Empirical evidence from real leaves under field conditions is scant and traditionally based on point measurements that do not capture spatial variation in heat load. We used thermal imagery under field conditions to measure the leaf thermal time constant (τ) in summer and the leaf‐to‐air temperature difference (?T) and temperature range across laminae (T_range) during winter, autumn and summer for 68 Proteaceae species. We investigated the influence of leaf area and margin complexity relative to effective leaf width (w_e), the latter being a more direct indicator of boundary layer thickness. Normalized difference of margin complexity had no or weak effects on thermal dynamics, but w_e strongly predicted τ and ?T, whereas leaf area influenced T_range. Unlike artificial leaves, however, spatial temperature distribution in large leaves appeared to be governed largely by structural variation. Therefore, we agree that small size, specifically w_e, has adaptive value in hot environments but not with the idea that thermal regulation is the primary evolutionary driver of leaf dissection.     

'Photoinhibition' of Heliconia under natural tropical conditions: the importance of leaf orientation for light interception and leaf temperature

The influence of irradiance on photosynthesis under natural conditions was studied in aseasonal Singapore using three Heliconia taxa: H. rostrata, H. psittacorum × H. spathocircinata cv. Golden Torch and H. psittacorum cv. Tay. When grown under full sunlight, all three heliconias exhibited reduced phatosynthetic capacities and lowered chlorophyll content per leaf area as compared with those grown under intermediate and deep shade. A marked decrease in the chlorophyll fluorescence F_v/Fm ratio and an increase in photochemical quenching (1- q_p) and non-photochemical quenching (q_N) were observed in upper leaves of plants grown under full sunlight. Increases in q_N suggest that ‘photoinhibition’ (decreases in F_v/F_m) in Heliconia grown under natural tropical conditions are probably due to photoprotective energy dissipation processes. The quantum yield, the maximum photosynthetic rate, F_v/F_m and the chlorophyll content of upper leaves were lower than those of lower leaves on the same plants grown under full sunlight. Similarly, lower values were obtained for the tip (sun) portion than for the base (shaded) portion of the leaves. The changes in F_v/F_m and in the levels of (1 –q_p) in leaves grown under intermediate and deep shade were negligible in plants during the course of day. However, there was a steep decrease in F_v/F_m and an increase in the levels of (1 –q_p), along with an increase in incident light in the sun leaves. The lowest F_v/F_m and the highest level of (1 –q_p) indicated minimum PSII efficiency at midday in full sun. These results indicate that, in Heliconia, the top leaves (particularly leaf tips) experienced sustained decreases in PSII efficiency upon exposure to full sunlight. Although all three taxa exhibited sustained decreases in photosynthetic capacity in full sunlight, the sun leaves of ‘Tay’ showed higher photosynthetic capacity than those of the other two taxa. This could be due, at least in part, to the vertical leaf angle and smaller lamina area. When the upright leaves of ‘Tay’ were constrained to a horizontal angle, they exhibited lower PSII efficiency (F_vIF_m ratio), while horizontal leaves of ‘Rostrata’ and ‘Golden Torch’ inclined lo near-vertical angles showed increased efficiency. Thus, an increase in leaf angle helps to achieve a reduction in the sustained decrease in PSII efficiency by decreasing the levels of incident sunlight and subsequently the leaf temperature.     

Spatial variability of above‐ground net primary production in Uruguayan grasslands: a remote sensing approach

Question: How does above‐ground net primary production (ANPP) differ (estimated from remotely sensed data) among vegetation units in sub‐humid temperate grasslands? Location: Centre‐north Uruguay. Methods: A vegetation map of the study area was generated from LANDSAT imagery and the landscape configuration described. The functional heterogeneity of mapping units was analysed in terms of the fraction of photosynthetically active radiation absorbed by green vegetation (fPAR), calculated from the normalized difference vegetation index (NDVI) images provided by the moderate resolution imaging spectroradiometer (MODIS) sensor. Finally, the ANPP of each grassland class was estimated using NDVI and climatic data. Results: Supervised classification presented a good overall accuracy and moderate to good average accuracy for grassland classes. Meso‐xerophytic grasslands occupied 45% of the area, Meso‐hydrophytic grasslands 43% and Lithophytic steppes 6%. The landscape was shaped by a matrix of large, unfragmented patches of Meso‐xerophytic and Meso‐hydrophytic grasslands. The region presented the lowest anthropic fragmentation degree reported for the Rio de la Plata grasslands. All grassland units showed bimodal annual fPAR seasonality, with spring and autumn peaks. Meso‐hydrophytic grasslands showed a radiation interception 10% higher than the other units. On an annual basis, Meso‐hydrophytic grasslands produced 3800 kg dry matter (DM) ha^?1 yr^?1 and Meso‐xerophytic grasslands and Lithophytic steppes around 3400 kg·DM·ha^?1·yr^?1. Meso‐xerophytic grasslands had the largest spatial variation during most of the year. The ANPP temporal variation was higher than the fPAR variability. Conclusions: Our results provide valuable information for grazing management (identifying spatial and temporal variations of ANPP) and grassland conservation (identifying the spatial distribution of vegetation units).     

Effects of continuous leaf wetness on photosynthesis: adverse aspects of rainfall

Above-ground parts of Phaseolus vulgaris L. plants were treated with artificial misty rain (‘rain’) in a growth chamber to investigate the effects of leaf wetness on photosynthetic performance. The following results were obtained. (1) Stomata closed completely within 2 min of the onset of continuous ‘rain’ application and gradually opened to half the original aperture by 60 min. The rate of CO₂ exchange measured on such wet leaves changed in parallel with the changes in stomatal aperture and attained 60 to 70% of the control level by 1h. (2) The dependence of the rate of leaf photosynthesis, A, on the intercellular CO₂ concentration, c_i [A(c_i) relationship], examined in thoroughly dried leaves which had been treated with ‘rain’ did not change until after 4 h of treatment. However, leaves treated for 6h showed discernible decreases in A at high c_i (c_i>500μmolmol ^?1). The photosynthetic rate of leaves treated with ‘rain’ for 24 h was reduced at all c_i, and A at the ambient CO₂ concentration of 350μmolmol^?1 was 60 to 70% of that of the control level. The rate of photosynthesis did not recover even after 3 d of treatment of the plants in a dry environment. These results clearly indicate that leaf wetness causes not only instantaneous suppression of photosynthesis but also chronic damage to the photosynthetic apparatus. Potential effects of leaf wetness on photosynthetic performance in nature are also discussed.     

Model simulations of spatial distributions and daily totals of photosynthesis in Eucalyptus grandis canopies

Summary A simulation model for radiation absorption and photosynthesis was used to test the hypothesis that observed nonuniform distributions of nitrogen concentrations in young Eucalyptus grandis trees result in greater amounts of daily assimilation than in hypothetical trees with uniform N distributions. Simulations were performed for trees aged 6, 9, 12 and 16 months which had been grown in plantations under a factorial combination of two levels of fertilization and irrigation. Observed leaf N distribution patterns yielded daily assimilation rates which were only marginally greater (<5%) than for hypothetical trees with uniform distributions. Patterns of assimilation distribution in individual tree crowns closely resembled those for absorbed radiation, rather than for N. These conclusions were unaffected by three choices of alternative leaf area density distributions. The simulation model was also used to calculate hourly and daily rates of canopy assimilation to investigate the relative importance of radiation absorption and total canopy nitrogen on assimilation. Simulated hourly rates of carbon assimilation were often lightsaturated, whereas daily carbon gain was directly proportional to radiation absorbed by the tree crown and to total mass of N in the leaves. Leaf nitrogen concentrations determined photosynthetic capacity, whereas total leaf area determined the amount of radiation absorbed and thus the degree to which capacity was realized. Observed total leaf area and total crown N were closely correlated. The model predicted that nitrogen use efficiences (NUE, mol CO₂ mol^–1 N) were 60% higher for unfertilized than for fertilized trees at low levels of absorbed photosynthetically active radiation (PAR). Nitrogen use efficiency was dependent on fertilizer treatment and on the amount of absorbed PAR; NUE declined with increasing absorbed PAR, but decreased more rapidly for unfertilized than for fertilized trees. Annual primary productivity was linearly related to both radiation absorbed and to mass of N in the canopy.     

Climate and plant trait strategies determine tree carbon allocation to leaves and mediate future forest productivity

Forest leaf area has enormous leverage on the carbon cycle because it mediates both forest productivity and resilience to climate extremes. Despite widespread evidence that trees are capable of adjusting to changes in environment across both space and time through modifying carbon allocation to leaves, many vegetation models use fixed carbon allocation schemes independent of environment, which introduces large uncertainties into predictions of future forest responses to atmospheric CO₂ fertilization and anthropogenic climate change. Here, we develop an optimization‐based model, whereby tree carbon allocation to leaves is an emergent property of environment and plant hydraulic traits. Using a combination of meta‐analysis, observational datasets, and model predictions, we find strong evidence that optimal hydraulic–carbon coupling explains observed patterns in leaf allocation across large environmental and CO₂ concentration gradients. Furthermore, testing the sensitivity of leaf allocation strategy to a diversity in hydraulic and economic spectrum physiological traits, we show that plant hydraulic traits in particular have an enormous impact on the global change response of forest leaf area. Our results provide a rigorous theoretical underpinning for improving carbon cycle predictions through advancing model predictions of leaf area, and underscore that tree‐level carbon allocation to leaves should be derived from first principles using mechanistic plant hydraulic processes in the next generation of vegetation models.     

Contributions of diffusional limitation, photoinhibition and photorespiration to midday depression of photosynthesis in Arisaema heterophyllum in natural high light

Diurnal changes in photosynthetic gas exchange and chlorophyll fluorescence were measured under full sunlight to reveal diffusional and non‐diffusional limitations to diurnal assimilation in leaves of Arisaema heterophyllum Blume plants grown either in a riparian forest understorey (shade leaves) or in an adjacent deforested open site (sun leaves). Midday depressions of assimilation rate (A) and leaf conductance of water vapour were remarkably deeper in shade leaves than in sun leaves. To evaluate the diffusional (i.e. stomatal and leaf internal) limitation to assimilation, we used an index [1–A/A₃₅₀], in which A₃₅₀ is A at a chloroplast CO₂ concentration of 350 μ mol mol ^{? 1}. A₃₅₀ was estimated from the electron transport rate (J_T), determined fluorometrically, and the specificity factor of Rubisco (S), determined by gas exchange techniques. In sun leaves under saturating light, the index obtained after the ‘peak’ of diurnal assimilation was 70% greater than that obtained before the ‘peak’, but in shade leaves, it was only 20% greater. The photochemical efficiency of photosystem II ( Δ F/F_m ′ ) and thus J_T was considerably lower in shade leaves than in sun leaves, especially after the ‘peak’. In shade leaves but not in sun leaves, A at a photosynthetically active photon flux density (PPFD) > 500 μ mol m ^{? 2} s ^{? 1} depended positively on J_T throughout the day. Electron flows used by the carboxylation and oxygenation (J_O) of RuBP were estimated from A and J_T. In sun leaves, the J_O/J_T ratio was significantly higher after the ‘peak’, but little difference was found in shade leaves. Photorespiratory CO₂ efflux in the absence of atmospheric CO₂ was about three times higher in sun leaves than in shade leaves. We attribute the midday depression of assimilation in sun leaves to the increased rate of photorespiration caused by stomatal closure, and that in shade leaves to severe photoinhibition. Thus, for sun leaves, increased capacities for photorespiration and non‐photochemical quenching are essential to avoid photoinhibitory damage and to tolerate high leaf temperatures and water stress under excess light. The increased Rubisco content in sun leaves, which has been recognized as raising photosynthetic assimilation capacity, also contributes to increase in the capacity for photorespiration.     

Photosynthetic capacity,irradiance and sequential senescence of sugar beet leaves

In field-grown sugar beet plants (Beta vulgaris L. cv. Dobrovická A), each of66 successive leaves produoed in the course of the vegetation period was different with respect to its photosynthetic capaoity (P_c), life span, duration of leaf area expansion, and longevity after its maximum leaf area (A_max) has developed. The proportionality between the seasonal changes in these characteristics was not the same if the sequential senescence of leaves was taken into account. With aging of individual leaves, P_c increased with the leaf area expansion having attained the peak value between 75% to 100% of A_max The rate of ontogenetic changes in P_c of each leaf was specified by the rate of its growth and development so that even at comparable ages the successive leaves constituted a series of different physiological units. The seasonal changes in quantum irradiance (PAR) were found to be responsible for differences in the growth characteristics between the successive leaves: Leaf expansion period was related with daily integrals of the incoming PAR (Io), while leaf longevity, after the A_max had been attained, was closely linked with PAR intercepted by the canopy (I). P_c expressed per the total leaf area of the plant was significantly correlated withI, while P_c calculated per unit leaf area of the plant was related toI _o Leaf potential to adapt P_c correspondingly to changes in PAR was greatest during leaf blade expansion; after the leaf had ceased to expand, changes in P_c were independent of differences in leaf irradiance. The results stress, at least for field conditions, the inadmissibility of the extrapolation of attributes from one leaf to the other ones sequentially senescing on the plant.     

Comparing PAR transmission models for forest understorey vegetation

Question: Do Beer's Law models, multi‐layer scattering models, and a semi‐empirical model for predicting PAR transmission through understorey vegetation give comparable results? Do different driving variables (LAI, PLAI and percentage cover) give different results? How do the models vary when fit with species‐specific, species‐average and the ‘default’ parameters recommended in the literature? Location: Upland boreal forests of western North America. Methods: In calibration and validation plots, PAR transmission was measured, total cover visually estimated, and leaf dispersion, PLAI and cover estimated for each species using a point‐frame. Leaf inclination was measured by clinometer. PAR transmission was modelled using empirically‐fit Beer's Law models, a semi‐empirical model based on hemispherical gap fraction and first‐order scattering, and a multi‐layer model allowing multiple scattering. All models were modified to use leaf area index (LAI), vertically projected leaf area index (PLAI), or percentage cover data. Results: The empirical Beer's Law models had the least bias and best precision in predicting PAR transmission. The semi‐empirical model also had little bias and good precision, since the scattering coefficient compensated for problems in the estimation of gap fraction. The multi‐layer model consistently underestimated transmission. There was little benefit in accounting for species separately. LAI and PLAI‐based models were the most precise, but percentage cover models also provided reasonable predictions of PAR transmission. Conclusions: PAR transmission through forest understories can be simply modelled with Beer's Law using one empirical coefficient representing the average understorey species. More complex scattering models are less effective, likely because they fail to account for the complexity of the dispersion of this vegetation layer and its effect on radiation scattering.     

Leaf nitrogen, photosynthesis, conductance and transpiration: scaling from leaves to canopies

A model is presented which solves simultaneously for leaf-scale stomatal conductance, CO₂ assimilation and the energy balance as a function of leaf position within canopies of well-watered vegetation. Fluxes and conductances were calculated separately for sunlit and shaded leaves. A linear dependence of photosynthetic capacity on leaf nitrogen content was assumed, while leaf nitrogen content and light intensity were assumed to decrease exponentially within canopies. Separate extinction coefficients were used for diffuse and direct beam radiation. An efficient Gaussian integration technique was used to compute fluxes and mean conductances for the canopy. The multilayer model synthesizes current knowledge of radiation penetration, leaf physiology and the physics of evaporation and provides insights into the response of whole canopies to multiple, interacting factors. The model was also used to explore sources of variation in the slopes of two simple parametric models (nitrogen- and light-use efficiency), and to set bounds on the magnitudes of the parameters. For canopies low in total N, daily assimilation rates are ～10% lower when leaf N is distributed uniformly than when the same total N is distributed according to the exponentially decreasing profile of absorbed radiation. However, gains are negligible for plants with high N concentrations. Canopy conductance, G_c should be calculated as G_c=Aσ(f_slg_sl+f_shg_sh), where Δ is leaf area index, f_si and f_sh are the fractions of sunlit and shaded leaves at each level, and g_si and g_sh are the corresponding stomatal conductances. Simple addition of conductances without this weighting causes errors in transpiration calculated using the ‘big-leaf’ version of the Penman-Monteith equation. Partitioning of available energy between sensible and latent heat is very responsive to the parameter describing the sensitivity of stomata to the atmospheric humidity deficit. This parameter also affects canopy conductance, but has a relatively small impact on canopy assimilation. Simple parametric models are useful for extrapolating understanding from small to large scales, but the complexity of real ecosystems is thus subsumed in unexplained variations in parameter values. Simulations with the multilayer model show that both nitrogen- and radiation-use efficiencies depend on plant nutritional status and the diffuse component of incident radiation, causing a 2- to 3-fold variation in these efficiencies.     

Seasonal variability in the effect of elevated CO2 on ecosystem leaf area index in a scrub-oak ecosystem

We report effects of elevated atmospheric CO₂ concentration (C_a) on leaf area index (LAI) of a Florida scrub‐oak ecosystem, which had regenerated after fire for between three and five years in open‐top chambers (OTCs) and was yet to reach canopy closure. LAI was measured using four nondestructive methods, calibrated and tested in experiments performed in calibration plots near the OTCs. The four methods were: PAR transmission through the canopy, normalized difference vegetation index (NDVI), hemispherical photography, and allometric relationships between plant stem diameter and plant leaf area. Calibration experiments showed: (1) Leaf area index could be accurately determined from either PAR transmission through the canopy or hemispherical photography. For LAI determined from PAR transmission through the canopy, ecosystem light extinction coefficient (k) varied with season and was best described as a function of PAR transmission through the canopy. (2) A negative exponential function described the relationship between NDVI and LAI; (3) Allometric relationships overestimated LAI. Throughout the two years of this study, LAI was always higher in elevated C_a, rising from, 20% during winter, to 55% during summer. This seasonality was driven by a more rapid development of leaf area during the spring and a relatively greater loss of leaf area during the winter, in elevated C_a. For this scrub‐oak ecosystem prior to canopy closure, increased leaf area was an indirect mechanism by which ecosystem C uptake and canopy N content were increased in elevated C_a. In addition, increased LAI decreased potential reductions in canopy transpiration from decreases in stomatal conductance in elevated C_a. These findings have important implications for biogeochemical cycles of C, N and H₂O in woody ecosystems regenerating from disturbance in elevated C_a.     

Estimating photosynthetic radiation use efficiency using incident light and photosynthesis of individual leaves

It has been theorized that photosynthetic radiation use efficiency (PhRUE) over the course of a day is constant for leaves throughout a canopy if leaf nitrogen content and photosynthetic properties are adapted to local light so that canopy photosynthesis over a day is optimized. To test this hypothesis, 'daily' photosynthesis of individual leaves of Solanum melongena plants was calculated from instantaneous rates of photosynthesis integrated over the daylight hours. Instantaneous photosynthesis was estimated from the photosynthetic responses to photosynthetically active radiation (PAR) and from the incident PAR measured on individual leaves during clear and overcast days. Plants were grown with either abundant or scarce N fertilization. Both net and gross daily photosynthesis of leaves were linearly related to daily incident PAR exposure of individual leaves, which implies constant PhRUE over a day throughout the canopy. The slope of these relationships (i.e. PhRUE) increased with N fertilization. When the relationship was calculated for hourly instead of daily periods, the regressions were curvilinear, implying that PhRUE changed with time of the day and incident radiation. Thus, linearity (i.e. constant PhRUE) was achieved only when data were integrated over the entire day. Using average PAR in place of instantaneous incident PAR increased the slope of the relationship between daily photosynthesis and incident PAR of individual leaves, and the regression became curvilinear. The slope of the relationship between daily gross photosynthesis and incident PAR of individual leaves increased for an overcast compared with a clear day, but the slope remained constant for net photosynthesis. This suggests that net PhRUE of all leaves (and thus of the whole canopy) may be constant when integrated over a day, not only when the incident PAR changes with depth in the canopy, but also when it varies on the same leaf owing to changes in daily incident PAR above the canopy. The slope of the relationship between daily net photosynthesis and incident PAR was also estimated from the photosynthetic light response curve of a leaf at the top of the canopy and from the incident PAR above the canopy, in place of that measured on individual leaves. The slope (i.e. net PhRUE) calculated in this simple way did not differ statistically from that calculated using data from individual leaves.