Adaptation of a PAM-fluorometer for remote sensing of chlorophyll fluorescence

The Laser-PAM described in this paper is an adaptation of the PAM 101 fluorometer (Heinz Walz, Effeltrich, Germany) designed for remote sensing and non-invasive laboratory measurements of chlorophyll fluorescence. It is based on a 5 mW laser diode, emitting at 638 nm, and a Fresnel lens coupled to the ED-101 detection unit. Due to these modifications, measurements can be performed at a distance ranging from 0.3 to 2 m. The ED-101 detection unit has been modified to perform simultaneous measurements of both modulated fluorescence and light reflected by the leaf. Reflected light showed a good estimation of the photosynthetically active radiation measured exactly at the same area as the fluorescence. A particular advantage of the Laser-PAM fluorometer is its suitability for remote measurements under field conditions. Simultaneous fluorescence and gas-exchange measurements, performed on grapevine leaves, are reported as an example of an application for the Laser-PAM. This revised version was published online in June 2006 with corrections to the Cover Date.     

Estimation of vegetation photosynthetic capacity from space‐based measurements of chlorophyll fluorescence for terrestrial biosphere models

Photosynthesis simulations by terrestrial biosphere models are usually based on the Farquhar's model, in which the maximum rate of carboxylation (V_cmax) is a key control parameter of photosynthetic capacity. Even though V_cmax is known to vary substantially in space and time in response to environmental controls, it is typically parameterized in models with tabulated values associated to plant functional types. Remote sensing can be used to produce a spatially continuous and temporally resolved view on photosynthetic efficiency, but traditional vegetation observations based on spectral reflectance lack a direct link to plant photochemical processes. Alternatively, recent space‐borne measurements of sun‐induced chlorophyll fluorescence (SIF) can offer an observational constraint on photosynthesis simulations. Here, we show that top‐of‐canopy SIF measurements from space are sensitive to V_cmax at the ecosystem level, and present an approach to invert V_cmax from SIF data. We use the Soil‐Canopy Observation of Photosynthesis and Energy (SCOPE) balance model to derive empirical relationships between seasonal V_cmax and SIF which are used to solve the inverse problem. We evaluate our V_cmax estimation method at six agricultural flux tower sites in the midwestern US using spaced‐based SIF retrievals. Our V_cmax estimates agree well with literature values for corn and soybean plants (average values of 37 and 101 μmol m^?2 s^?1, respectively) and show plausible seasonal patterns. The effect of the updated seasonally varying V_cmax parameterization on simulated gross primary productivity (GPP) is tested by comparing to simulations with fixed V_cmax values. Validation against flux tower observations demonstrate that simulations of GPP and light use efficiency improve significantly when our time‐resolved V_cmax estimates from SIF are used, with R² for GPP comparisons increasing from 0.85 to 0.93, and for light use efficiency from 0.44 to 0.83. Our results support the use of space‐based SIF data as a proxy for photosynthetic capacity and suggest the potential for global, time‐resolved estimates of V_cmax.     

实验课新方向：通过荧光反应探究影响光合作用的外界因素

光合作用是植物进行空气营养的一种方式,是绿色植物组织的基本功能,但是环境中的各种因素都会影响该过程。通过荧光光谱测定燕麦（Avenna sativa L,cv Prevision）植株的离体完整叶片的吸收光谱和作用光谱来探究影响光合作用的各种物理和化学因素,如高光强、高温、除草剂3,4-二氯苯-1,1-二甲基脲（DCMU）等。该实验方法适合植物生理学或生化专业的学生进行课外拓展研究。     

Spectral vegetation indices as the indicator of canopy photosynthetic productivity in a deciduous broadleaf forest

Aims Understanding of the ecophysiological dynamics of forest canopy photosynthesis and its spatial and temporal scaling is crucial for revealing ecological response to climate change. Combined observations and analyses of plant ecophysiology and optical remote sensing would enable us to achieve these studies. In order to examine the utility of spectral vegetation indices (VIs) for assessing ecosystem-level photosynthesis, we investigated the relationships between canopy-scale photosynthetic productivity and canopy spectral reflectance over seasons for 5 years in a cool, temperate deciduous broadleaf forest at 'Takayama' super site in central Japan.Methods Daily photosynthetic capacity was assessed by in situ canopy leaf area index (LAI), (LAI × V cmax [single-leaf photosynthetic capacity]), and the daily maximum rate of gross primary production (GPP max) was estimated by an ecosystem carbon cycle model. We examined five VIs: normalized difference vegetation index (NDVI), enhanced vegetation index (EVI), green–red vegetation index (GRVI), chlorophyll index (CI) and canopy chlorophyll index (CCI), which were obtained by the in situ measurements of canopy spectral reflectance.Important findings Our in situ observation of leaf and canopy characteristics, which were analyzed by an ecosystem carbon cycling model, revealed that their phenological changes are responsible for seasonal and interannual variations in canopy photosynthesis. Significant correlations were found between the five VIs and canopy photosynthetic capacity over the seasons and years; four of the VIs showed hysteresis-type relationships and only CCI showed rather linear relationship. Among the VIs examined, we applied EVI–GPP max relationship to EVI data obtained by Moderate Resolution Imaging Spectroradiometer to estimate the temporal and spatial variation in GPP max over central Japan. Our findings would improve the accuracy of satellite-based estimate of forest photosynthetic productivity in fine spatial and temporal resolutions, which are necessary for detecting any response of terrestrial ecosystem to meteorological fluctuations.     

Sun‐induced fluorescence – a new probe of photosynthesis: First maps from the imaging spectrometer HyPlant

Variations in photosynthesis still cause substantial uncertainties in predicting photosynthetic CO₂ uptake rates and monitoring plant stress. Changes in actual photosynthesis that are not related to greenness of vegetation are difficult to measure by reflectance based optical remote sensing techniques. Several activities are underway to evaluate the sun‐induced fluorescence signal on the ground and on a coarse spatial scale using space‐borne imaging spectrometers. Intermediate‐scale observations using airborne‐based imaging spectroscopy, which are critical to bridge the existing gap between small‐scale field studies and global observations, are still insufficient. Here we present the first validated maps of sun‐induced fluorescence in that critical, intermediate spatial resolution, employing the novel airborne imaging spectrometer HyPlant. HyPlant has an unprecedented spectral resolution, which allows for the first time quantifying sun‐induced fluorescence fluxes in physical units according to the Fraunhofer Line Depth Principle that exploits solar and atmospheric absorption bands. Maps of sun‐induced fluorescence show a large spatial variability between different vegetation types, which complement classical remote sensing approaches. Different crop types largely differ in emitting fluorescence that additionally changes within the seasonal cycle and thus may be related to the seasonal activation and deactivation of the photosynthetic machinery. We argue that sun‐induced fluorescence emission is related to two processes: (i) the total absorbed radiation by photosynthetically active chlorophyll; and (ii) the functional status of actual photosynthesis and vegetation stress.     

绿盲蝽为害对枣树叶片光合作用及叶绿素荧光特性的影响

【目的】探讨绿盲蝽Apolygus lucorum(Meyer-Dür)为害对枣树叶片光合作用的影响及其机制。【方法】以一年生冬枣Zizyphus jujuba cv.Dongzao和酸枣Ziziphus jujuba var.spinosa树叶片为试材,测定了绿盲蝽为害1,3,5和7 d时枣树叶片光合速率、气体交换、叶绿素荧光参数和叶绿素含量的变化。【结果】绿盲蝽为害3,5和7 d时冬枣叶片的净光合速率(net photosynthesis rate,Pn)较对照分别降低了55.83%,55.42%和59.61%;而酸枣叶片净光合速率仅在5和7 d时较对照分别降低了26.66%和27.34%。冬枣叶片的气孔导度被绿盲蝽为害3,5和7 d时较对照明显降低。冬枣叶片光合速率的下降与气孔导度(stomatal conductance,Gs)和总叶绿素含量的下降呈显著正相关,而酸枣叶片光合速率的下降仅与叶绿素含量显著正相关。绿盲蝽为害后冬枣和酸枣叶片的快速荧光诱导曲线受到显著影响。冬枣叶片的最大光化学效率(maximum photochemical efficiency,Fv/Fm)在绿盲蝽为害不同时间时相对于对照明显降低,而酸枣叶片没有受到明显的影响。绿盲蝽为害不同时间对冬枣和酸枣叶片的光系统Ⅱ放氧复合体(oxygen-evolving complex,OEC)以及光反应活性中心均造成了伤害,但酸枣受到的伤害程度明显低于冬枣。绿盲蝽为害5和7 d后冬枣叶片的光系统Ⅱ的电子传递活性降低,而酸枣叶片光系统Ⅱ的电子传递活性没有受到显著影响。绿盲蝽为害导致冬枣和酸枣叶片的电子传递的量子产额较对照明显降低,酸枣叶片中的降低幅度低于冬枣。【结论】绿盲蝽为害造成枣树叶片净光合速率明显降低,不同品种存在明显差异,冬枣叶片Pn降低程度明显高于酸枣。绿盲蝽为害后枣树叶片净光合速率的下降与叶绿素含量降低呈显著正相关。绿盲蝽为害影响了枣树叶片PSⅡ的结构和功能,导致供体侧的OEC受到伤害,光合作用PSⅡ反应中心失活,PSⅡ反应中心关闭程度增加,电子传递活性受到了抑制,其中酸枣叶片PSⅡ受到的影响明显低于冬枣叶片。     

脱落酸对低温下雷公藤幼苗光合作用及叶绿素荧光的影响

以1年生雷公藤扦插苗为试材,研究低温胁迫下不同浓度外源脱落酸（ABA,0、5、10、15、20、25 mg·L^-1)叶面喷施处理对雷公藤叶片光合作用及叶绿素荧光参数的影响.结果表明：喷施20 mg·L^-1的ABA能显著提高雷公藤幼苗的抗冷性,减缓低温下雷公藤叶片净光合速率(P_n)、蒸腾速率(T_r)、气孔导度(g_s)、胞间CO₂浓度(C_i)的下降幅度,提高幼苗叶片的光合能力.低温处理6 d后,随着ABA浓度上升,雷公藤叶片的初始荧光(F_o)下降,最大荧光(F_m)和PSII最大光化学效率(F_v/F_m)上升,PSII实际光化学量子产量(Φ_PSⅡ)、光化学猝灭系数(q_P)先下降后上升,而非光化学猝灭系数(q_N)呈下降－上升－下降趋势.P_n、g_s、q_P、F_m和F_v/F_m均在20 mg·L^-1ABA处理时达到峰值.不同浓度ABA的相对电子传递速率(rETR)随着光化光强度增加呈先上升后下降的趋势,当光化光强度(PAR)达到395 μmol·m^-2s^-1时,各处理的rETR达到最高值,其中25 mg·L^-1和20 mg·L^-1ABA处理分别比对照高17.1%和5.2%.雷公藤叶片Φ_PSⅡ的光响应曲线均随光化光强度升高而下降,q_N的光响应曲线则呈相反趋势.     

Observation and characterisation of a transient in the yield of chlorophyll fluorescence in intact spinach chloroplasts

A transient in chlorophyll fluorescence, which is associated with a transient in 9-aminoacridine fluorescence and a perturbation in the rate of oxygen evolution, has been observed in intact spinach chloroplasts. The results indicate that changes in the redox state of Q are, at least partially, responsible for the transient in chlorophyll fluorescence. The size of the transient is highly dependent upon the concentration of inorganic phosphate and upon the pH of the medium. The properties of the transient are consistent with the suggestion that it reflects changes in the levels of stromal intermediates during induction.Abbreviations BES NN-Bis(2-hydroxyethyl)2-aminoethanesulphonic acid dihydroxyacetone-P(DHAP): dihydroxyacetone phosphate glycerate-3-P (PGA): glycerate-3-phosphate - HEPES N-2-Hydroxyethylpiperazine-N-2-ethanesulphonic acid - MES 2-(N-Morpholino)ethanesulphonic acid - Pi inorganic phosphate - qE quenching of chlorophyll fluorescence by the energisation of the thylakoid membrane - qQ quenching of chlorophyll fluorescence by oxidised Q, the electron acceptor of photosystem 2 - ribose-5-P (R5P) ribose-5-phosphate - Rbu-5-P ribulose-5-phosphate     

内生固氮菌对甘蔗伸长期光合生理特性的影响

利用联合生物固氮是有效降低甘蔗生产成本的途径之一.在温室无氮栽培条件下,将内生固氮菌株A01分别接种到6个不同基因型甘蔗(GT24、GT9、YC84/153、GT18、CP65/357和F172)中,在甘蔗伸长期测定+1叶片的叶绿素含量(叶绿素a、叶绿素b)、光合作用参数净光合速率(Pn)、胞间CO2浓度(Ci)、气孔...     

C3 photosynthesis in Aristida longifolia: Implication for photosynthetic diversification in Aristidoideae (Poaceae)

Only a small percentage of plant species undergo C(4) photosynthesis. Despite its rarity, the C(4) pathway has evolved numerous times from C(3) ancestors, with as many as 18 independent origins in grasses alone. We report non-Kranz (C(3)) anatomy in Aristida longifolia, a species in a genus of ca. 300 species previously thought to possess only Kranz (C(4)) anatomy. Leaf blade transections of A. longifolia show widely spaced vascular bundles, nonradiate chlorenchyma, and few or no chloroplasts in cells of the sheaths surrounding the vascular bundle, all features indicative of C(3) photosynthesis. Carbon isotope ratios range from -27.68 to -29.71%, likewise indicative of C(3) photosynthesis. We also reconstruct the phylogeny of Aristidoideae, comprising Aristida, Sartidia (C(3)), and Stipagrostis (C(4)), using a sample of 11 species, including A. longifolia, and DNA sequences of the nuclear ribosomal internal transcribed spacer region and the chloroplast rpl16 intron and trnL-trnF region. Sartidia and Stipagrostis resolve as sisters, and sister to this clade is Aristida. Aristida longifolia resolves as sister to the remaining species in the genus. C(3) photosynthesis is hypothesized to be ancestral in Aristidoideae, which means the C(4) pathway evolved twice in the subfamily-in Stipagrostis and early in the diversification of the Aristida clade.     

Chlorophyll a fluorescence: can it shed light on fundamental questions in photosynthetic carbon dioxide fixation?

Abstract A close, immediate and precise relationship between chlorophyll a fluorescence and photosynthetic carbon assimilation in vivo is demonstrated. The examples discussed include kinetics displayed during dark to light transitions plus oscillations and transients observed during changes in the gas phase surrounding the leaf. Remaining uncertainties surrounding the relationship between chlorophyll fluorescence and photosynthesis are attributed to the underlying complexity of the regulatory mechanisms involved. Examples are also given that show how multiple simultaneous measurements of different aspects of the photosynthetic process may contribute to the resolution of these uncertainties. The practical relevance of these matters is also discussed, particularly in relation to the limitations of the photosynthetic process and to the use of chlorophyll fluorescence as a diagnostic probe of chemical and genetic manipulation and stress.     

和田设施温室6个葡萄品种光合和 叶绿素荧光日变化特征

【目的】探讨和田地区设施温室葡萄品种光合和叶绿素荧光日变化特征,综合评价其光合能力强弱,为该地区温室葡萄引种及栽培管理措施制定提供参考依据。【方法】以和田设施温室引入6个葡萄品种‘户太八号’‘妮娜女皇’‘夏黑’‘阳光玫瑰’‘新郁’‘克瑞森无核’为试验材料,测定并分析各品种光合有效辐射（PAR）、叶绿素相对含量（SPAD）、光合与叶绿素荧光参数,并利用主成分分析对各鲜食葡萄品种能力进行综合性评价。【结果】（1）设施温室光合有效辐射在不同位置总体表现出棚前＞棚后＞棚中,在不同架面位置总体表现为架上＞架中＞架下。（2）葡萄叶片SPAD值在不同架面位置表现为架上部＞架中部＞架下部,在品种间由高到低依次为‘克瑞森无核’‘夏黑’‘新郁’‘户太八号’‘阳光玫瑰’‘妮娜女皇’。（3）各品种Pn、Gs、Tr均呈现出双峰曲线的日变化趋势,Ci日变化总体呈现U形或W形变化规律;Fv/Fo与Fv/Fm总体上呈现出先下降后上升的日变化规律。（4）6个葡萄品种的光合能力表现为‘克瑞森无核’＞‘夏黑’＞‘阳光玫瑰’＞‘新郁’＞‘户太八号’＞‘妮娜女皇’。【结论】 ‘克瑞森无核’与‘夏黑’相比其他品种有较高的Pn、Gs、Tr、 Fo、Fm和较低的Fv/Fo与Fv/Fm,对和田地区适应性较强,能够适应当地的高温与高光强设施环境。     

Red fluorescence spectra as a new means of determining the rate of damage to photosynthesis apparatus in plants caused by stress

A new method is presented for determining the rate of damage to photosynthesis apparatus efficiency caused by stress using the red fluorescence spectra of plant leaves. A direct connection was found between the position of the red fluorescence maximum _max and the photosynthesis apparatus efficiency . The method was tested on several examples and good results were obtained.     

Chlorophyll fluorescence transients of Photosystem II membrane particles as a tool for studying photosynthetic oxygen evolution

The rise of the chlorophyll fluorescence yield of Photosystem II (PS II) membranes as induced by high-intensity actinic light comprises only two distinct phases: (1) the initial O-J increase and (2) the subsequent J-P increase. Partial inhibition of the PS II donor side by heating or washing procedures which remove peripheral PS II proteins or cofactors of the oxygen-evolving complex results in decrease of magnitude and rate of the J-P phase. The rate constant of the J-P increase is directly proportional to the steady-state rate of oxygen evolution; complete suppression of the J-P phase corresponds to full inhibition. A characteristic dip after J-level is observed only in Tris-washed or severely heated PS II membranes; manganese release correlates with appearance of the dip after J-level as verified by EPR spectroscopy. Presence of stabilizing cosolutes (glycine betaine, sucrose) or addition of donor-side cofactors (bicarbonate, chloride, calcium) to PS II membranes before heating (47 °C, 5 min) diminishes J-P phase suppression and prevents dip appearance, whereas the addition after heating is without effect. In conclusion, analysis of chlorophyll fluorescence transients of PS II membranes is a potentially useful tool for investigations on photosynthetic oxygen evolution. A decreased rate of the J-P phase can be employed as a convenient indicator for partial inhibition of oxygen-evolution activity; the appearance of a dip after J-level is suggestive of manganese release. This revised version was published online in June 2006 with corrections to the Cover Date.     

热带雨林下砂仁叶片光合作用和叶绿素荧光参数在雾凉季和雨季的日变化

于雾凉季和雨季研究了自然条件下透光率不同的3块雨林样地下砂仁叶片光合参数和叶绿素荧光参数的日变化。结果表明：(1)砂仁叶片净光合速率(Pn)与叶表光量子通量密度(PFD)和气孔导度(Gs)呈正相关,与胞间CO2浓度呈负相关,PFD是影响Pn的主导因素。雾凉季砂仁叶片Pn和Gs及其日变幅均低于雨季,相同光强下雾凉季Pn及表观光能和CO2利用效率也低于雨季,这与雾凉季的光强和温度较低有关。(2)随日间光强的增加光系统Ⅱ最大光能转换效率(Fv／Fm)、光系统Ⅱ光能捕获效率(Fv‘／Fm’)和电子传递量子效率(ΦPSI)下降,非光化学猝灭系数(NPQ)和电子传递速率则上升,黄昏各参数都能恢复到黎明时的水平。表明随光强的增加,砂仁热耗散增多,日间光抑制加重,这种光抑制是光合功能下调的保护性反应而非光破坏。(3)Fv／Fm、Fv‘／Fm’和ΦPSI与PFD呈负相关,NPQ与PFD呈正相关。相同光强下雾凉季Fv／Fm、Fv’／Fm’和ΦPSI低于、NPQ则高于雨季,这与雾凉季Pn较低有关。(4)随栽培时间的延长砂仁衰老和产量降低均不明显,砂仁的产量随生长环境光强的增加而呈升高趋势。各季节不同光强下生长的砂仁在午间高光强时均未发生光合机构的破坏,且随光强的升高其Pn呈上升趋势,说明砂仁具有适应更高光强并获得高产的潜力。     

Remote sensing of heterogeneity in photosynthetic efficiency, electron transport and dissipation of excess light in Populus deltoides stands under ambient and elevated CO2 concentrations, and in a tropical forest canopy, using a new laser-induced fluorescence transient device

Determining the spatial and temporal diversity of photosynthetic processes in forest canopies presents a challenge to the evaluation of biological feedbacks needed for improvement of carbon and climate models. Limited access with portable instrumentation, especially in the outer canopy, makes remote sensing of these processes a priority in experimental ecosystem and climate change research. Here, we describe the application of a new, active, chlorophyll fluorescence measurement system for remote sensing of light use efficiency, based on analysis of laser‐induced fluorescence transients (LIFT). We used mature stands of Populus grown at ambient (380 ppm) and elevated CO₂ (1220 ppm) in the enclosed agriforests of the Biosphere 2 Laboratory (B2L) to compare parameters of photosynthetic efficiency, photosynthetic electron transport, and dissipation of excess light measured by LIFT and by standard on‐the‐leaf saturating flash methods using a commercially available pulse‐modulated chlorophyll fluorescence instrument (Mini‐PAM). We also used LIFT to observe the diel courses of these parameters in leaves of two tropical forest dominants, Inga and Pterocarpus, growing in the enclosed model tropical forest of B2L. Midcanopy leaves of both trees showed the expected relationships among chlorophyll fluorescence‐derived photosynthetic parameters in response to sun exposure, but, unusually, both displayed an afternoon increase in nonphotochemical quenching in the shade, which was ascribed to reversible inhibition of photosynthesis at high leaf temperatures in the enclosed canopy. Inga generally showed higher rates of photosynthetic electron transport, but greater afternoon reduction in photosynthetic efficiency. The potential for estimation of the contribution of outer canopy photosynthesis to forest CO₂ assimilation, and assessment of its response to environmental stress using remote sensing devices such as LIFT, is briefly discussed.     

不同底质对海菜花叶绿素荧光诱导动力学参数及净光合速率的影响

海菜花通常被认为是对富营养化水体较为敏感的沉水植物,但近年来研究认为其对富营养化水体也有较好的净化作用。目前,海菜花在滇池回植取得一定进展,但回植时能否直接利用富营养化湖泊底泥为底质或是需要添加养分含量较少的土壤改良底泥未有明确的结论。通过对比不同底质上海菜花的光合生理参数表明:种植40d时,在湖沙和大理洱海底泥上种植的海菜花光合能力最好,红壤上种植的海菜花净光合速率仅为洱海底泥上种植海菜花的63.6%,其最大荧光(Fm)和可变荧光(Fv)比洱海底泥上种植的海菜花分别降低52.5%和58.8%,反应中心捕获的用于电子传递的能量(Eto/RC)仅为洱海底泥上种植海菜花的59.7%,光系统Ⅱ(PSⅡ)最大光化学量子产量(Fv/Fm)、性能指数(PI_(ABS))和反应中心数量(RC/CSo)也最低,叶片的单位反应中心耗散的能量(DIo/RC)是洱海底泥上种植海菜花的2.48倍,表明在红壤上种植的海菜花叶片的光合能力较低。种植80d时,不同底质上种植的海菜花的光合生理指标发生了一定改变,其中红壤上种植的海菜花光合能力大幅提升,净光合速率和性能指数等均有提高,与洱海底泥上种植的海菜花已无明显差异;然而红壤上种植的海菜花虽然在80d时的光合性能恢复,但其叶片长度已受到影响,明显低于其他底质上的海菜花的叶片长度。与此同时湖沙上种植的海菜花的光合能力在其他底质上种植的海菜花均有所增加的情况下明显下降,其净光合速率明显低于其他底质上种植的海菜花,单位面积的活性反应中心数量显著降低。结合分析不同底质的理化性质,在种植初期红壤呈酸性可能是导致海菜花叶片光合能力低的原因,之后红壤长期处于厌氧环境使得土壤pH升高后光合能力恢复正常,但其叶片的长度在短期内未有明显增加,表明其光合产物的累积受到了影响;而湖沙上种植的海菜花后期光合能力减弱可能是由于后期水体中养分耗尽导致的。因此,在回植海菜花过程中,可直接利用富营养化湖泊中的底泥,养分含量丰富的底泥不会成为其生长的限制因素,而酸性土壤可能还会限制其生长。     

蕾后期和花前期切花菊品种'神马'不同部位叶片光合作用和叶绿素荧光特性的比较

对蕾后期和花前期切花菊(Chrysanthemum morifolium Ramat.)品种'神马'('Jinba')不同部位叶片光合作用参数日变化、叶绿素荧光参数、光响应曲线及参数进行了研究.结果表明:蕾后期和花前期品种'神马'叶片蒸腾速率(Tr)、气孔导度(Gs)和净光合速率(Pn)的日变化均为单峰曲线,峰值出现在10:00或12:00;胞间CO2浓度(Ci)的日变化则先降低后升高,谷值出现在12:00.蕾后期和花前期品种'神马'叶片Tr、Ci和Gs值的平均值总体上随叶片位置降低而逐渐升高;蕾后期不同部位叶片Pn值的平均值差异较小,花前期叶片Pn值的平均值则随叶片位置降低而逐渐降低.随着叶片位置降低,蕾后期和花前期品种'神马'叶片的初始荧光(Fo)、最大荧光(Fm)、可变荧光(Fv)、表观量子效率(AQE)和最大净光合速率(Pmax)以及蕾后期的暗呼吸速率(Rd)均逐渐降低,而花前期的Rd值以及蕾后期和花前期的光补偿点(LCP)均逐渐升高.随着光合有效辐射(PAR)升高,蕾后期不同部位叶片以及花前期中部叶和下部叶的Pn值呈先急剧升高后趋于平稳的变化趋势,而花前期上部叶的Pn值则呈先急剧升高后逐渐下降的变化趋势.研究结果显示:在切花菊设施栽培过程中适当补充光照可提高切花菊品质.     

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.