Importance of the green color,absorption gradient,and spectral absorption of chloroplasts for the radiative energy balance of leaves

Terrestrial green plants absorb photosynthetically active radiation (PAR; 400–700 nm) but do not absorb photons evenly across the PAR waveband. The spectral absorbance of photosystems and chloroplasts is lowest for green light, which occurs within the highest irradiance waveband of direct solar radiation. We demonstrate a close relationship between this phenomenon and the safe and efficient utilization of direct solar radiation in simple biophysiological models. The effects of spectral absorptance on the photon and irradiance absorption processes are evaluated using the spectra of direct and diffuse solar radiation. The radiation absorption of a leaf arises as a consequence of the absorption of chloroplasts. The photon absorption of chloroplasts is strongly dependent on the distribution of pigment concentrations and their absorbance spectra. While chloroplast movements in response to light are important mechanisms controlling PAR absorption, they are not effective for green light because chloroplasts have the lowest spectral absorptance in the waveband. With the development of palisade tissue, the incident photons per total palisade cell surface area and the absorbed photons per chloroplast decrease. The spectral absorbance of carotenoids is effective in eliminating shortwave PAR (<520 nm), which contains much of the surplus energy that is not used for photosynthesis and is dissipated as heat. The PAR absorptance of a whole leaf shows no substantial difference based on the spectra of direct or diffuse solar radiation. However, most of the near infrared radiation is unabsorbed and heat stress is greatly reduced. The incident solar radiation is too strong to be utilized for photosynthesis under the current CO₂ concentration in the terrestrial environment. Therefore, the photon absorption of a whole leaf is efficiently regulated by photosynthetic pigments with low spectral absorptance in the highest irradiance waveband and through a combination of pigment density distribution and leaf anatomical structures.     

Photocontrol of the Functional Coupling between Photosynthesis and Stomatal Conductance in the Intact Leaf : Blue Light and Par-Dependent Photosystems in Guard Cells

The photocontrol of the functional coupling between photosynthesis and stomatal conductance in the leaf was investigated in gas exchange experiments using monochromatic light provided by lasers. Net photosynthesis and stomatal conductance were measured in attached leaves of Malva parviflora L. as a function of photon irradiance at 457.9 and 640.0 nanometers.

Photosynthetic rates and quantum yields of photosynthesis were higher under red light than under blue, on an absorbed or incident basis.

Stomatal conductance was higher under blue than under red light at all intensities. Based on a calculated apparent photon efficiency of conductance, blue and red light had similar effects on conductance at intensities higher than 0.02 millimoles per square meter per second, but blue light was several-fold more efficient at very low photon irradiances. Red light had no effect on conductance at photon irradiances below 0.02 millimoles per square meter per second. These observations support the hypothesis that stomatal conductance is modulated by two photosystems: a blue light-dependent one, driving stomatal opening at low light intensities and a photosynthetically active radiation (PAR)-dependent one operating at higher irradiances.

When low intensity blue light was used to illuminate a leaf already irradiated with high intensity, 640 nanometers light, the leaf exhibited substantial increases in stomatal conductance. Net photosynthesis changed only slightly. Additional far-red light increased net photosynthesis without affecting stomatal conductance. These observations indicate that under conditions where the PAR-dependent system is driven by high intensity red light, the blue light-dependent system has an additive effect on stomatal conductance.

The wavelength dependence of photosynthesis and stomatal conductance demonstrates that these processes are not obligatorily coupled and can be controlled by light, independent of prevailing levels of intercellular CO₂. The blue light-dependent system in the guard cells may function as a specific light sensor while the PAR-dependent system supplies a CO₂-modulated energy source providing functional coupling between the guard cells and the photosynthesizing mesophyll.

     

大豆叶片光合、蒸腾的日变化及其与生态因子的相关性研究

两年的试验结果表明：大豆叶片光合作用、蒸腾作用、叶水势及叶片对水分扩散的导性在大田自然条件中呈“单峰”或“双峰”型日变化。其变化的产生并非系生物内生物节奏引起,而实属同期生态条件影响的结果。在本研究中,光强、叶温及水蒸汽压亏缺是主要诱因。这些生态因子在量上的不同组合,导致所测生理特性的日变化呈不同趋势。     

Rapid stomatal responses to humidity

The response of leaf conductance in apple to rapid changes in atmospheric humidity was studied using a continuous flow porometer. Leaf-air vapour pressure difference was changed by adjusting the humidity of the inlet air or by altering the flow rate of the air through the chamber. The time course of the response of leaf conductance to leaf-air vapour pressure difference was monitored for periods up to 10 min using a chart-recorder. There were significant changes in leaf conductance within seconds of changing humidity. These were attributed to alterations in stomatal aperture.Abbreviations E evaporation rate - g leaf conductance - PAR photosynthetically active radiation     

曼陀罗光合特性研究

采用LI-6400型便携式光合测定仪对潜在能源植物曼陀罗(Datura stramonium L.)在6月份至9月份的光合特性指标(净光合速率、光合有效辐射强度和气孔导度)的日变化以及净光合速率和叶绿素含量的月平均值变化进行了研究,并采用非直角双曲线对曼陀罗叶片净光合速率的光响应曲线和C02响应曲线进行了拟合.结果表明:实验期间(6月份至9月份),光合有效辐射强度和曼陀罗叶片净光合速率的日变化均呈"单峰型",峰值出现在11:00左右;7月份的净光合速率峰值最高,为19.01μmol·m-2·s-1;气孔导度日变化呈"双峰型",最大峰值和次峰值分别出现在12:00和14:00.不同月份的净光合速率月平均值有极显著差异(P＜0.01),其中7月份的净光合速率月平均值最大,为9.41 μmol·m-2·s-1;不同月份叶绿素的月平均含量变化与净光合速率月平均值的变化相似,7月份叶绿素的月平均含量也最高,但叶绿素a、b及总叶绿素月平均含量有一定差异.曼陀罗叶片净光合速率的光响应曲线和CO2响应曲线相似,在光合有效辐射强度和CO2浓度较低的条件下呈线性升高,至饱和点后缓慢升高并趋于稳定;曼陀罗叶片的光补偿点为22.42 μmol·m-2·s-1,光饱和点为689.26 μmol·m-2·s-1;CO2补偿点为74.06μmol·mol-1,CO2饱和点为1 331.97μmol·mol-1.研究结果说明,曼陀罗为具有一定耐阴能力的阳性植物,但CO2同化能力较弱.     

Energy exchange within individual layers of a meadow

Summary Measurements of the radiation extinction in a meadow at Baumkirchen (Tyrol) show that the decrease in the photosynthetically active radiation (PhAR: 400–700 nm) is different to that of the total net-radiation in a characteristic way. The photosynthetically active radiation is distributed evenly to all vegetation layers, the active surface comprises practically the entire 90 cm high canopy. The total radiation energy (net-radiation) is absorbed and 45% is converted into sensible and latent heat only in a 25 cm wide layer, i.e., between 30 and 55 cm within the canopy. A second active surface lies in the lowermost 10 cm of the meadow and at the soil surface where additional 28% of the radiation energy is transformed into heat.     

海岸带复合农林业系统植物种群光环境特征研究

本文对三种银杏-农作物复合模式下植物种群对光合有效辐射的削弱和截获进行了模型化分析,并探讨了不同模式下光照强度的时空分布规律。结果表明:银杏果用-叶用-豆类作物复合模式具有最好的复合光效益,光能截获率可达92%;PAR在植冠层的削弱遵循Beer-Lambert定律;植物种群的叶面积、地上部分生物量、光吸收的关系可以用Y=ax^b数学模型描述。同时,果用、材用银杏冠层有较大的透光性和光强变异系数,可以作为银杏-农作物初期经营的上层树种,但要注意冠形的调控;光强的时间变化受冠层条件和太阳高度角的双重影响。     

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)。     

Mechanisms of amplification of photosynthetically active radiation in the symbiotic deep-water coral Leptoseris fragilis

The symbiotic coral Leptoseris fragilis lives in the Red Sea at depths of 95–145 m. Symbiotic dinoflagellates (zooxanthellae) themselves possess well known adaptations to low light intensities. In L. fragilis we found indications that light amplifying mechanisms of the host improve photosynthesis of the symbionts. Light of short wavelengths is absorbed by host pigments which transform short into longer wavelengths. The transformed light is more efficient for photosynthesis. Action spectra measurements of photosynthesis demonstrated the amplification of photosynthetically active radiation. Monochromatic light of 387 nm (outside the main absorption maxima of the algal pigments) at subsaturation photon flux densities was as effective photosynthetically as polychromatic light of 415–490 nm, which fits the absorption maxima of the zooxanthellae.     

Phototropins promote plant growth in response to blue light in low light environments

Phototropins (phot1 and phot2) are plant-specific blue light receptors for phototropism, chloroplast movement, leaf expansion, and stomatal opening. All these responses are thought to optimize photosynthesis by helping to capture light energy efficiently, reduce photodamage, and acquire CO2. However, experimental evidence for the promotion of plant growth through phototropins is lacking. Here, we report dramatic phototropin-dependent effects on plant growth. When plants of Arabidopsis thaliana wild type, the phot1 and phot2 mutants, and the phot1 phot2 double mutant were grown under red light, no significant growth differences were observed. However, if a very low intensity of blue light (0.1 micromol m(-2) s(-1)) was superimposed on red light, large increases in fresh weight up to threefold were found in those plants that carried functional PHOT1 genes. When the intensity of blue light was increased to 1 micromol m(-2) s(-1), the growth enhancement was also found in the phot1 single mutant, but not in the double mutant, indicating that phot2 mediated similar responses as phot1 with a lower sensitivity. The effects occurred under low photosynthetically active radiation in particular. The well-known physiological phototropin-mediated responses, including chloroplast movement, stomatal opening, and leaf expansion, in the different lines tested indicated an involvement of these responses in the blue light-induced growth enhancement. We conclude that phototropins promote plant growth by controlling and integrating a variety of responses that optimize photosynthetic performance under low photosynthetically active radiation in the natural environment.     

Combined impacts of irradiance and dehydration on leaf hydraulic conductance: insights into vulnerability and stomatal control

The leaf is a hydraulic bottleneck, accounting for a large part of plant resistance. Thus, the leaf hydraulic conductance (K(leaf) ) is of key importance in determining stomatal conductance (g(s) ) and rates of gas exchange. Previous studies showed that K(leaf) is dynamic with leaf water status and irradiance. For four species, we tested the combined impacts of these factors on K(leaf) and on g(s) . We determined responses of K(leaf) and g(s) to declining leaf water potential (Ψ(leaf) ) under low and high irradiance (<6 and >900 μmol photons m(-2) s(-1) photosynthetically active radiation, respectively). We hypothesized greater K(leaf) vulnerability under high irradiance. We also hypothesized that K(leaf) and g(s) would be similar in their responses to either light or dehydration: similar light-responses of K(leaf) and g(s) would stabilize Ψ(leaf) across irradiances for leaves transpiring at a given vapour pressure deficit, and similar dehydration responses would arise from the control of stomata by Ψ(leaf) or a correlated signal. For all four species, the K(leaf) light response declined from full hydration to turgor loss point. The K(leaf) and g(s) differed strongly in their light- and dehydration responses, supporting optimization of hydraulic transport across irradiances, and semi-independent, flexible regulation of liquid and vapour phase water transport with leaf water status.     

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.     

Inversion of net ecosystem CO2 flux measurements for estimation of canopy PAR absorption

The fractional absorption of photosynthetically active radiation (f_PAR) is frequently a key variable in models describing terrestrial ecosystem–atmosphere interactions, carbon uptake, growth and biogeochemistry. We present a novel approach to the estimation of the fraction of incident photosynthetically active radiation absorbed by the photosynthetic components of a plant canopy (f_Chl). The method uses micrometeorological measurements of CO₂ flux and incident radiation to estimate light response parameters from which canopy structure is deduced. Data from two Ameriflux sites in Oklahoma, a tallgrass prairie site and a wheat site, are used to derive 7‐day moving average estimates of f_Chl during three years (1997–1999). The inverse estimates are compared to long‐term field measurements of PAR absorption. Good correlations are obtained when the field‐measured f_PAR is scaled by an estimate of the green fraction of total leaf area, although the inverse technique tends to be lower in value than the field measurements. The inverse estimates of f_Chl using CO₂ flux measurements are different from measurements of f_PAR that might be made by other, more direct, techniques. However, because the inverse estimates are based on observed canopy CO₂ uptake, they might be considered more biologically relevant than direct measurements that are affected by non‐physiologically active components of the canopy. With the increasing number of eddy covariance sites around the world the technique provides the opportunity to examine seasonal and inter‐annual variation in canopy structure and light harvesting capacity at individual sites. Furthermore, the inverse f_Chl provide a new source of data for development and testing of f_PAR retrieval using remote sensing. New remote sensing algorithms, or adjustments to existing algorithms, might thus become better conditioned to ‘biologically significant’ light absorption than currently possible.     

Light Absorption and Competition in Mixed Communities of Hordeum Leporinum-Euphorbia Scordifolia

The effect of Euphorbia scordifolia and Hordeum leporinum competition on leaf area development, radiant energy absorption, and dry matter production was evaluated in a field experiment. Profile measurements (0-0.3, 0.3-0.6, 0.6-0.9, and >0.9 m above ground) of absorbed photosynthetically active radiation (APAR) and leaf area index (LAI) by species were taken at four densities of E. scordifolia (0, 1, 4, and 12 plants per m²). APAR calculated for H. leporinum in mixed communities was 79, 77, and 49 % of the APAR in H. leporinum and LAI was reduced to 81, 65, and 37 %. LAI of H. leporinum was concentrated in the 0.3-0.6 m layer, while the taller E. scordifolia plants had the greatest LAI above 0.6 m. By absorbing radiant energy in the upper canopy, E. scordifolia reduced APAR penetrating to H. leporinum. Measurements of net photosynthetic and transpiration rates, leaf temperature, and stomatal conductance confirmed the importance of competition for PAR for plant growth and metabolism.     

An integrated portable apparatus for the simultaneous field measurement of photosynthetic CO2 and water vapour exchange, light absorption and chlorophyll fluorescence emission of attached leaves

Abstract. A portable apparatus has been constructed to measure simultaneously the quantum yield of CO₂ assimilation, light absorption, chlorophyll fluorescence emission and water vapour exchange of attached intact leaves in the field. The core of the instrument is a light-integrating spherical leaf chamber which includes ports for a light source, photosynthetically active radiation sensor, fluorescence probes and gas inlet and outlet manifolds. Measurement of the quantum flux inside the empty chamber and with a leaf present allows determination of leaf absorptance. An open gas exchange system is employed using an infra-red analyser to measure leaf CO₂ exchange. Using a DC white light source the quantum yield of CO₂ assimilation based on absorbed light (φ_abs) may be determined rapidly in either ambient air or artificial gas mixtures. Inclusion of capacitance humidity probes into the gas inlet and outlet ports allows simultaneous determination of water vapour exchange and subsequent estimation of stomatal conductance to CO₂ and intercellular CO₂ concentration. Measurement of fluorescence emission by the sample leaf exposed to white light is achieved by a modulated fluorescence detection system. In addition to determination of the minimal, maximal and variable fluorescence levels, a further analysis allows the photochemical and non-photochemical components of fluorescence quenching, to be estimated. The theory and design of this apparatus is described in detail. The use of the apparatus in the field is demonstrated through a study of the photosynthetic performance of a maize and bean crop during the growing season and by analysis of the photosynthetic performance of crops subjected to nitrogen-stress and a herbicide treatment.     

Diel oscillation in the optical activity of carotenoids in the absorption spectrum of <Emphasis Type="Italic">Nannochloropsis</Emphasis>

In this paper we show that the absorption spectrum of the microalgae Nannochloropsis oceanica exhibits changes in response to the modulation of incident light. A model was used to analyze the contribution of different active pigments to the total absorption in the photosynthetically active radiation region and suggested consistent diel oscillations in the optical activity of carotenoids.     

Comparing global models of terrestrial net primary productivity (NPP): importance of vegetation structure on seasonal NPP estimates

Estimates of the seasonal absorbed fraction of photosynthetically active radiation (FPAR) and net primary productivity (NPP) are compared among four production efficiency models (PEMs) and seven terrestrial biosphere models simulating canopy development. In addition, the simulated FPARs of the models are compared to the FASIR-FPAR derived from NOAA-AVHRR satellite observations. All models reproduce observed summergreen phenology of temperate deciduous forests rather well, but perform less well for raingreen phenology of savannas. Some models estimate a much longer active canopy in savannas than indicated by satellite observations. As a result, these models estimate high negative monthly NPP during the dry season. For boreal and tropical evergreen ecosystems, several models overestimate LAI and FPAR. When the simulated canopy does respond to unfavourable periods, the seasonal NPP is largely determined by absorbed photosynthetically active radiation (APAR). When the simulated canopy does not respond to unfavourable periods, the light use efficiency (LUE) influences the seasonal NPP more. However, the relative importance of APAR and LUE can change seasonally.     

Simultaneous measurement of stomatal conductance, non-photochemical quenching, and photochemical yield of photosystem II in intact leaves by thermal and chlorophyll fluorescence imaging

A new imaging system capable of simultaneously measuring stomatal conductance and fluorescence parameters, non-photochemical quenching (NPQ) and photochemical yield of photosystem II (Phi(PSII)), in intact leaves under aerobic conditions by both thermal imaging and chlorophyll fluorescence imaging was developed. Changes in distributions of stomatal conductance and fluorescence parameters across Phaseolus vulgaris L. leaves induced by abscisic acid treatment were analyzed. A decrease in stomatal conductance expanded in all directions from the treatment site, then mainly spread along the lateral vein toward the leaf edge, depending on the ABA concentration gradient and the transpiration stream. The relationships between stomatal conductance and fluorescence parameters depended on the actinic light intensity, i.e. NPQ was greater and Phi(PSII) was lower at high light intensity. The fluorescence parameters did not change, regardless of stomatal closure levels at a photosynthetically active photon flux (PPF) of 270 micro mol m(-2) s(-1); however, they drastically changed at PPF values of 350 and 700 micro mol m(-2) s(-1), when the total stomatal conductance decreased to less than 80 and 200 mmol m(-2) s(-1), respectively. This study has, for the first time, quantitatively analyzed relationships between spatiotemporal variations in stomatal conductance and fluorescence parameters in intact leaves under aerobic conditions.     

Light Absorption and Competition in Mixed Communities of Hordeum Leporinum-Euphorbia Scordifolia

The effect of Euphorbia scordifolia and Hordeum leporinum competition on leaf area development, radiant energy absorption, and dry matter production was evaluated in a field experiment. Profile measurements (0-0.3, 0.3-0.6, 0.6-0.9, and >0.9 m above ground) of absorbed photosynthetically active radiation (APAR) and leaf area index (LAI) by species were taken at four densities of E. scordifolia (0, 1, 4, and 12 plants per m²). APAR calculated for H. leporinum in mixed communities was 79, 77, and 49 % of the APAR in H. leporinum and LAI was reduced to 81, 65, and 37 %. LAI of H. leporinum was concentrated in the 0.3-0.6 m layer, while the taller E. scordifolia plants had the greatest LAI above 0.6 m. By absorbing radiant energy in the upper canopy, E. scordifolia reduced APAR penetrating to H. leporinum. Measurements of net photosynthetic and transpiration rates, leaf temperature, and stomatal conductance confirmed the importance of competition for PAR for plant growth and metabolism. This revised version was published online in June 2006 with corrections to the Cover Date.     

Light acclimation during and after leaf expansion in soybean

Soybean plants (Glycine max var. Ransom) were grown at light intensities of 850 and 250 μeinsteins m⁻² sec⁻¹ of photosynthetically active radiation. A group of plants was shifted from each environment into the other environment 24 hours before the beginning of the experiment. Net photosynthetic rates and stomatal conductances were measured at 2,000 and 100 μeinsteins m⁻² sec⁻¹ photosynthetically active radiation on the 1st, 2nd, and 5th days of the experiment to determine the time course of photosynthetic light adaptation. The following factors were also measured: dark respiration, leaf water potential, leaf thickness, internal surface area per external surface area, chlorophyll content, photosynthetic unit size and number, specific leaf weight, and activities of malate dehydrogenase, and glycolate oxidase. Comparisons were made with plants maintained in either 850 or 250 μeinsteins m⁻² sec⁻¹ environments. Changes in photosynthesis, stomatal conductance, leaf anatomy, leaf water potential, photosynthetic unit size, and glycolate oxidase activity occurred upon altering the light environment, and were complete within 1 day, whereas chlorophyll content, numbers of photosynthetic units, specific leaf weight, and malate dehydrogenase activity showed slower changes. Differences in photosynthetic rates at high light were largely accounted for by internal surface area differences with low environmental light associated with low internal area and low photosynthetic rate. An exception to this was the fact that plants grown at 250 μeinsteins m⁻² sec⁻¹ then switched to 850 μeinsteins m⁻² sec⁻¹ showed lower photosynthesis at high light than any other treatment. This was associated with higher glycolate oxidase and malate dehydrogenase activity. Photosynthesis at low light was higher in plants kept at or switched to the lower light environment. This increased rate was associated with larger photosynthetic unit size, and lower dark respiration and malate dehydrogenase activity. Both anatomical and physiological changes with environmental light occurred even after leaf expansion was complete and both were important in determining photosynthetic response to light.