Photosynthetic characteristics and photosynthesis-light response curve models of summer maize

Aim The photosynthesis-light response curve is the most commonly used method to explore the relationship between photosynthetically active radiation and the net photosynthetic rate, because it is more effective to reflect the plant photosynthetic characteristics. And it is very meaningful for researchers to choose a suitable summer corn (Zea mays) photosynthetic model and optimal the models of photosynthesis-light response curve by analyzing the differences between simulation and observation in each growth period of some plant. So the object of this paper was to propose some useful suggestions for the choice of summer corn photosynthetic modes and the optimization of the photosynthetic light response curves model in further.Methods In this paper, three typical photosynthetic models were used to fitting the photosynthetic light response curve for upper leaf, leaf at ear of grain and lower leaf of summer corn during bell and milk period. And then the fitting degree of each model was compared to the measured data. Photosynthetic active radiation was divided into three parts, and the fitting residual errors of these three models were analyzed individually.Important findings The photosynthetic characteristic parameters such as maximum net photosynthetic rate (P_nmax), saturated light intensity (I_sat) and dark respiration rate (R_d) decreased constantly with a top-down leaf position and the parameters at milk stage were generally lower than the bell stage. Each growth period and leaf position could fit the curve, but some deviation exists for the P_nmax and I_sat in the rectangular hyperbolic model and the non-rectangular hyperbolic model. The results of curve fitting residual showed that the simulation values from Ye Zi-Piao model were closest to the actual values, and especially for the high photosynthetically active radiation section.     

夏玉米光合特性及光响应曲线拟合

通过对光合-光响应曲线的研究来探索光合有效辐射与净光合速率的关系是一种非常重要的手段。合适的模型才能较好地反映植株的光合特性。分析由于生育期与叶位的不同而导致各模型拟合值与实测值差异的变化性以及不同光合有效辐射强度下各模型的适用性, 可为夏玉米(Zea mays)光合模型的选择和光合-光响应曲线模型的进一步优化提供参考。该研究运用3种典型的光合模型对夏玉米大喇叭口期与乳熟期上部叶、穗位叶与下部叶做光合-光响应曲线拟合, 对比各模型的拟合度以及对实测数据的反映情况, 并将光合有效辐射分为3段, 分析3种模型在每段的拟合残差。结果表明: 最大净光合速率(P_nmax)、饱和光强(I_sat)、暗呼吸速率(R_d)等光合特征参数值随叶位由上而下呈降低趋势, 乳熟期普遍小于大喇叭口期; 不同生育期和叶位3种模型均可以拟合, 直角双曲线模型、非直角双曲线模型对P_nmax、I_sat的拟合值与实测值有一定的偏差; 通过残差分析表明叶子飘模型的拟合值与实测值最为相符, 尤其对高光部分的拟合表现出独有的优越性。     

人工长白落叶松冠层光合作用-光响应曲线最优模型

以黑龙江省帽儿山林场15年生人工长白落叶松为研究对象,采用直角双曲线模型(RH)、非直角双曲线模型(NRH)、指数模型(EM)、修正直角双曲线模型(MRH)和修正指数模型(MEM)分别对4种不同光响应特征的光合作用-光响应曲线(光抑制型光响应曲线,PLC_i;光饱和型光响应曲线,PLC_s;未饱和型光响应曲线,PLC_u;弱光环境下植被的光响应曲线,PLC_w)进行拟合,计算出光饱和时的最大净光合速率(P_{n max})、暗呼吸速率(R_d)、光补偿点(LCP)、光饱和点(LSP)及表观量子效率(AQY)等重要的光合生理指标,综合对比5个候选模型对不同响应曲线的拟合优度和对光合生理指标的估计精度.结果表明: MEM模型仅适用于拟合光抑制型曲线,MRH对光抑制型曲线和光饱和型曲线的拟合效果最好(R_a²分别为0.9986和0.9978),NRH最适合拟合未饱和型曲线和弱光环境型曲线(R_a²分别为0.9996和0.9963).在所有类型曲线中,MRH模型估计P_{n max}时,平均相对误差绝对值(MAPE)最低(0.1%R_d表现出更准确的估计(MAPE分别为1.8%、0.1%和3.9%);RH模型对光抑制型曲线的LCP及光饱和型曲线的R_d有更好的估计效果(MAPE分别为1.0%和2.7%);EM模型适用于估计弱光环境型曲线的LCP(MAPE为0.2%).MRH在保证较好的模型拟合效果及光合生理指标估计精度以外,还在拟合不同类型曲线时表现出极高的稳定性,因此,本文选择MRH模型作为拟合人工长白落叶松冠层光合作用-光响应曲线的最优模型.     

A comparison of the photosynthesis — light intensity relationship in phylogenetically different marine microalgae

Measured under equivalent physiological conditions, the photosynthesis-light intensity relationship based on oxygen production/mg chlorophyll a was found to be the same in five species representing the different chlorophyll c containing divisions of marine phytoplankton. Other non-photochemical metabolic processes related to photosynthesis such as diurnal variations, maximal photosynthesis rates, and dark oxygen uptake were quite different, and so these are the more significant factors in production and ecological distribution of diatoms, dinoflagellates, and coccolithophores. In contrast, the green algae tested showed a significantly different photosynthesis-light intensity curve from the chlorophyll c group.     

Changes in Net O(2) Exchange Induced by Inorganic Nitrogen in the Blue-Green Alga Anacystis nidulans

The response of net O₂ exchange to light intensity by intact Anacystis nidulans cells in the presence of saturating NaHCO₃ concentrations followed a curve with an inflection near the light-compensation point. Addition of either KNO₃ or NH₄Cl stimulated O₂ uptake in the dark and at light intensities below the light-compensation point. This resulted in steeper slopes of the curve calculated below and above the light-compensation point. At O₂ concentrations limiting dark respiration, addition of inorganic nitrogen had no effect on either dark respiration or O₂ exchange in the light. The apparent changes in photosynthetic yield observed under normal O₂ concentration disappeared when respiration was limited by O₂ availability, indicating that the effects of inorganic nitrogen on O₂ exchange at low light intensities are due to stimulation of respiration rather than to increases in photosynthetic yield.     

Exposure times in rapid light curves affect photosynthetic parameters in algae

Short-and long-duration light curves were applied to four macroalgae (Ulva sp., Codium fragile, Ecklonia radiata and Lessonia variegata), and two microalgal species (Chlorella emersonii and Chaetoceros muellerii). With increasing light curve duration, the maximal relative electron transport rate increased by a factor of three in E. radiata, and by factors of 1.25 and 1.23 in C. emersonii and L. variegata, respectively, but did not change in C. fragile and Ch. muellerii. The light saturation point E_k increased by 26 μmol photons m⁻² s⁻¹ in C. emersonii and 20 μmol photons m⁻² s⁻¹ in Ch. muellerii and E. radiata with elevated light curve exposure times. Oscillatory patterns of the continuous fluorescence readings reflect accumulation of Q_A. Continuous fluorescence values increased, or decreased, by approximately 10% within light curve increments. However, oscillations of 25% were not uncommon, which shows that cells are changing their photo-physiological response state during steady light conditions. Increasing dark acclimation times prior to light curve application lowered maximal relative electron transport rates in the C. emersonii (from 28 ± 1.7 to 25 ± 1.2 for 15 and 95 min dark acclimation in short-duration light curves respectively). This effect was counterbalanced by longer light curve application. It can therefore be concluded that manipulation of light exposure and dark incubation prior to the experiment affects the photosynthetic response, presumably due to different activation states of photosynthetic and photoprotective mechanisms. The highly species-specific photo-response patterns imply that a common rapid light curve protocol will generate artefacts in some species.     

Modeling study on photosynthetic-light response curves of a C4 plant,maize

Aims A light response curve can reflect a plant’s ability to utilize light, which is also a key tool in determining the relationship between photosynthetic capacity and environmental factors; however the model accuracies concerning the light response curve remain elusive. The objectives of this study were to compare and assess the model accuracies related to a light response curve and the effects of drought. Methods A field rain shelter was used to control the soil water conditions. To obtain photosynthesis parameters from the light response curve and the drought effects, the relevant models (including the rectangular model, non-rectangular hyperbolic model, modified rectangular hyperbolic model, exponential model, quadratic function model, and a newly modified model) were applied to fit the light response curves. The validity of each model was tested by analyzing the differences between the fitted values obtained by the models and the measured values. Important findings The newly modified model has been proved to performing relatively better in accurately describing the light response curve patterns, and credibly obtaining the crucial photosynthetic parameters such as the maximum net photosynthetic rate, light saturation point, light compensation point, and dark respiration rate, especially under high radiation conditions.     

Thermal acclimation of photosynthesis: on the importance of adjusting our definitions and accounting for thermal acclimation of respiration

While interest in photosynthetic thermal acclimation has been stimulated by climate warming, comparing results across studies requires consistent terminology. We identify five types of photosynthetic adjustments in warming experiments: photosynthesis as measured at the high growth temperature, the growth temperature, and the thermal optimum; the photosynthetic thermal optimum; and leaf-level photosynthetic capacity. Adjustments of any one of these variables need not mean a concurrent adjustment in others, which may resolve apparently contradictory results in papers using different indicators of photosynthetic acclimation. We argue that photosynthetic thermal acclimation (i.e., that benefits a plant in its new growth environment) should include adjustments of both the photosynthetic thermal optimum (T _opt) and photosynthetic rates at the growth temperature (A _growth), a combination termed constructive adjustment. However, many species show reduced photosynthesis when grown at elevated temperatures, despite adjustment of some photosynthetic variables, a phenomenon we term detractive adjustment. An analysis of 70 studies on 103 species shows that adjustment of T _opt and A _growth are more common than adjustment of other photosynthetic variables, but only half of the data demonstrate constructive adjustment. No systematic differences in these patterns were found between different plant functional groups. We also discuss the importance of thermal acclimation of respiration for net photosynthesis measurements, as respiratory temperature acclimation can generate apparent acclimation of photosynthetic processes, even if photosynthesis is unaltered. We show that while dark respiration is often used to estimate light respiration, the ratio of light to dark respiration shifts in a non-predictable manner with a change in leaf temperature.     

Change in Photosynthetic Capacity over the Cell Cycle in Light/Dark-Synchronized Amphidinium carteri Is Due Solely to the Photocycle

Cell cycle dependent photosynthesis in the marine dinoflagellate Amphidinium carteri was studied under constant illumination and light/dark (L/D) photocycles to distinguish intrinsic cell cycle control from environmental influences. Cells were grown in constant light and on a 14:10 L:D cycle at light intensities that would yield a population growth rate of 1 doubling per day. In the former case division was asynchronous, and cells were separated according to cell cycle stage using centrifugal elutriation. Cells grown on the L:D cycle were synchronized, with division restricted to the dark period. Cell cycle stage distributions were quantified by flow cytometry. Various cell age groups from the two populations were compared as to their photosynthetic response (photosynthetic rate versus irradiance) to determine whether or not the response was modulated primarily by cell cycle constraints or the periodic L/D cycle. Cell cycle variation in photosynthetic capacity was found to be determined solely by the L/D cycle; it was not present in cells grown in constant light.     

Seagrass Canopy Photosynthetic Response Is a Function of Canopy Density and Light Environment: A Model for Amphibolis griffithii

A three-dimensional computer model of canopies of the seagrass Amphibolis griffithii was used to investigate the consequences of variations in canopy structure and benthic light environment on leaf-level photosynthetic saturation state. The model was constructed using empirical data of plant morphometrics from a previously conducted shading experiment and validated well to in-situ data on light attenuation in canopies of different densities. Using published values of the leaf-level saturating irradiance for photosynthesis, results show that the interaction of canopy density and canopy-scale photosynthetic response is complex and non-linear, due to the combination of self-shading and the non-linearity of photosynthesis versus irradiance (P-I) curves near saturating irradiance. Therefore studies of light limitation in seagrasses should consider variation in canopy structure and density. Based on empirical work, we propose a number of possible measures for canopy scale photosynthetic response that can be plotted to yield isoclines in the space of canopy density and light environment. These plots can be used to interpret the significance of canopy changes induced as a response to decreases in the benthic light environment: in some cases canopy thinning can lead to an equivalent leaf level light environment, in others physiological changes may also be required but these alone may be inadequate for canopy survival. By providing insight to these processes the methods developed here could be a valuable management tool for seagrass conservation during dredging or other coastal developments.     

玉米全生育期黄化材料的光合性能分析

通过对玉米(Zea mays L.)黄化材料和2份正常自交系的光合特性、光响应曲线及荧光动力学参数进行分析,结果表明,黄化材料yglm1的净光合速率(Pn)、气孔导度(Gs)和胞间CO2浓度(Ci)均显著低于正常自交系Mo17和RP128,而蒸腾速率(Tr)却显著高于正常自交系;黄化材料的光补偿点、光饱和点、暗呼吸速率及表观量子效率均高于正常自交系Mo17和RP128,但其净光合速率却低于正常自交系;黄化材料的荧光动力学参数初始荧光(Fo)、最大荧光(Fm)、PSⅡ原初光能转换效率(Fv/Fm)、PSⅡ原初光能捕获效率(Fv'/Fm')、PSⅡ光下实际光化学效率(ΦPSⅡ)、光化学猝灭系数(qP)均显著低于正常自交系Mo17和RP128,非光化学猝灭系数(qN)显著高于正常自交系。相对于正常自交系,黄化材料具有较低的电子传递潜力、原初光能捕获效率和转换效率、光能分配及利用率,这可能是导致其净光合能力降低的根本原因。     

Physiological performances of the blue-green alga Anacystis nidulans in continuous turbidostat fermenter culture

A sterile continuous turbidostat culture in a 2-1 fermenter was used to systematically measure the gas exchange rates of Anacystis nidulans in a highly turbulent system under strictly controlled environmental conditions. An extensive physiological characterization of Anacystis is given in terms of photosynthesis rates (CO₂ uptake and O₂ evolution) and dark respiration rates as function of different parameters such as stirrer speed, temperature, CO₂ and O₂ concentration, light intensity, culture density and pH. Steady state ATP levels and apparent photophosphorylation rates complete the performance data. The dependence of the photosynthetic quotient from the parameters enables a physiological characterization of the light dependent nitrate assimilation.     

A coral polyp model of photosynthesis, respiration and calcification incorporating a transcellular ion transport mechanism

A numerical simulation model of coral polyp photosynthesis, respiration and calcification was developed. The model is constructed with three components (ambient seawater, coelenteron and calcifying fluid), and incorporates photosynthesis, respiration and calcification processes with transcellular ion transport by Ca-ATPase activity and passive transmembrane CO₂ transport and diffusion. The model calculates dissolved inorganic carbon and total alkalinity in the ambient seawater, coelenteron and calcifying fluid, dissolved oxygen (DO) in the seawater and coelenteron and stored organic carbon (CH₂O). To reconstruct the drastic variation between light and dark respiration, respiration rate dependency on DO in the coelenteron is incorporated. The calcification rate depends on the aragonite saturation state in the calcifying fluid (Ωa _cal). Our simulation result was a good approximation of “light-enhanced calcification.” In our model, the mechanism is expressed as follows: (1) DO in the coelenteron is increased by photosynthesis, (2) respiration is stimulated by increased DO in the light (or respiration is limited by DO depletion in the dark), then (3) calcification increases due to Ca-ATPase, which is driven by the energy generated by respiration. The model simulation results were effective in reproducing the basic responses of the internal CO₂ system and DO. The daily calcification rate, the gross photosynthetic rate and the respiration rate under a high-flow condition increased compared to those under the zero-flow condition, but the net photosynthetic rate decreased. The calculated calcification rate responses to variations in the ambient aragonite saturation state (Ωa _amb) were nonlinear, and the responses agreed with experimental results of previous studies. Our model predicted that in response to ocean acidification (1) coral calcification will decrease, but will remain at a higher value until Ωa _amb decreases to 1, by maintaining a higher Ωa _cal due to the transcellular ion transport mechanism and (2) the net photosynthetic rate will increase.     

Adaptation to CO(2) Level and Changes in the Phosphorylation of Thylakoid Proteins during the Cell Cycle of Chlamydomonas reinhardtii

The photosynthetic performance of synchronously grown Chlamydomonas reinhardtii alternated rhythmically during the cell cycle. The activity of the “CO₂ concentrating mechanism” including the ability to accumulate CO₂ internally and the activity of carbonic anhydrase peaked after 6 to 9 hours of light and reached minimum after 6 to 9 hours of dark. Consequently, the apparent photosynthetic affinity to extracellular CO₂ alternated rhythmically. At the end of the dark period the cells behaved as if they were adapted to high CO₂ even though they were continuously aerated with air. Results from experiments in which the light or dark periods were extended bear on the interaction between the internal (cell cycle or biological clock) and the external (light) signal. The observed rhythmical alterations in photosynthetic V_max may result from changes in PSII activity. The latter may be partly explained by the capacity for phosphorylation of thylakoid proteins, which reached maximum after 9 hours of light and decreased toward the dark period.     

A mechanistic model for the light response of photosynthetic electron transport rate based on light harvesting properties of photosynthetic pigment molecules

Models describing the light response of photosynthetic electron transport rate (ETR) are routinely used to determine how light absorption influences energy, reducing power and yields of primary productivity; however, no single model is currently able to provide insight into the fundamental processes that implicitly govern the variability of light absorption. Here we present development and application of a new mechanistic model of ETR for photosystem II based on the light harvesting (absorption and transfer to the core ‘reaction centres’) characteristics of photosynthetic pigment molecules. Within this model a series of equations are used to describe novel biophysical and biochemical characteristics of photosynthetic pigment molecules and in turn light harvesting; specifically, the eigen-absorption cross-section and the minimum average lifetime of photosynthetic pigment molecules in the excited state, which describe the ability of light absorption of photosynthetic pigment molecules and retention time of excitons in the excited state but are difficult to be measured directly. We applied this model to a series of previously collected fluorescence data and demonstrated that our model described well the light response curves of ETR, regardless of whether dynamic down-regulation of PSII occurs, for a range of photosynthetic organisms (Abies alba, Picea abies, Pinus mugo and Emiliania huxleyi). Inherent estimated parameters (e.g. maximum ETR and the saturation irradiance) by our model are in very close agreement with the measured data. Overall, our mechanistic model potentially provides novel insights into the regulation of ETR by light harvesting properties as well as dynamical down-regulation of PSII.     

Reversible pH Changes in Cells of Chlamydomonas reinhardi Resulting from CO(2) Fixation in the Light and Its Evolution in the Dark

Illumination of a suspension of Chlamydomonas reinhardi causes an increase in the pH of the medium which is reversed in the dark. This pH change is a manifestation of CO₂ uptake in the light and its evolution in the dark. Simultaneous measurements of pH changes and oxygen evolution reveal that the photosynthetic coefficient approaches one.     

Photosynthetic response of marine phytoplankton to eserine salicylate,a zooplankton anaesthetizing substance

Addition of eserine salicylate, a synaptic metabolic inhibitor, to marine algal cultures of Sketetonema costatum (Greville) Cleve, Dunaliella tertiolecta Butcher, and Prorocentrum micans Ehrenberg had a varied effect on the rate of respiration; α (the initial slope of the light saturation curve) and P^B_m (the saturation curve) and P (the specific production rate at optimal light intensity). As these effects were either concentration dependent, species specific or both, we find that it is not possible to manipulate photosynthesis-light relationships in a predictable manner by eserine salicylate treatment.     

Changes in photosynthesis caused by adaptation of maize seedlings to short-term drought

Detached leaf is in the state of increasing water deficit; it is a good experimental model for looking into the hardening effect of adaptation of eight-day-old maize (Zea mays L.) seedlings to short-term drought (five days without watering). The light stage of photosynthesis and photosynthetic CO₂/H₂O exchange in detached leaves were studied. Specific surface density of leaf tissue (SSDL), the content of chlorophylls a and b, proline, MDA as well as photosynthetic parameters: quantum yield of photosystem II fluorescence, assimilation of CO₂, and transpiration at room temperature and light saturation (density of PAR quantum flux of 2000 μmol/(m² s)) at normal and half atmospheric CO₂ concentration were determined. The leaves of seedlings exposed to short-term drought differed from control material by a greater SSDL and higher content of proline. The hardening effect of the stress agent on the dark stage of photosynthesis was detected; it was expressed in the maintenance of the higher photosynthetic CO₂ assimilation against control material due to the elevation of stomatal conductance for CO₂ diffusing into the leaf. Judging from the lack of differences in the MDA content, short-term drought did not injure photosynthetic membranes. In detached leaves of experimental maize seedlings, photosynthesis was maintained on a higher level than in control material.     

A mechanistic model of the adaptation of phytoplankton photosynthesis

The mechanistic model of the phytoplankton photosynthesis-light intensity relationship by Eilers and Peeters (1988.Ecol. Modelling 42, 199–215) is investigated mathematically. The model is based on the physiological idealization of transition probabilities between states of the photosynthetic factories,PSF. The model was found to have under constant light condition a globally stable unique positive equilibrium, while under periodically varying light (e.g. daily periodicity) there exists a unique globally asymptotically stable periodic solution. Based on this, the adaptation to a change of light intensity is defined as a process by which the state ofPSF converges to an equilibrium. Assuming that phytoplankton regulates its photosynthetic production rate with a certain strategy which maximizes production, two such possible strategies were examined. Both the instantaneous and the integral maximal photosynthetic production were shown to have the same result. With realistic qualitative assumptions of the shape of the dependence of the four model parameters on the light intensity to which phytoplankton is adapted, the numerical values of parameters under both constant and periodically varying conditions are determined by applying Pontryagin's maximum principle.