Different Relationship Between Electron Transport and CO2 Assimilation in two Zea mays Cultivars as Influenced by Increasing Irradiance

Gas exchange and fluorescence parameters were measured simultaneously in two Zea mays L. cultivars (Liri and 121C D8) to assess the relationship between the quantum yield of electron transport (_PS2) and the quantum yield of CO₂ assimilation (_CO2) in response to photosynthetic photon flux density (PPFD). The cv. Liri was grown under controlled environmental conditions in a climate chamber (CC) while cv. 121C D8 was grown in CC as well as outdoors (OT). By exposing the two maize cultivars grown in CC to an increasing PPFD, higher photosynthetic and photochemical rates were evidenced in cv. Liri than in cv. 121C D8. In Liri plants the _PS2/_CO2 ratio increased progressively up to 27 with increasing PPFD. This suggests that the reductive power was more utilised in non-assimilatory processes than in CO₂ assimilation at high PPFD. On the contrary, by exposing 121C D8 plants to increasing PPFD, _PS2/_CO2 was fairly constant (around 11–13), indicating that the electron transport rate was tightly down regulated by CO₂ assimilation. Although no significant differences were found between _PS2/_CO2 of the 121C D8 maize grown under CC and OT by exposing them to high PPFD, the photosynthetic rate and photochemical rates were higher in OT maize plants.     

Photosynthetic Responses of Tropical Trees to Short-Term Exposure to Ozone

Saplings of the tropical trees Tibouchina pulchra (Cham.) Cogn., Caesalpinia echinata Lam., and Psidium guajava L. cv. Paluma were exposed in open-top chambers with charcoal filtered air and measurements of gas exchange and chlorophyll fluorescence were made before (t₁) and after exposure to non-filtered air plus O₃ (t₂), simulating 6-h peaks of O₃ similar to those observed in São Paulo city (SE Brazil, reaching an AOT40 of 641 nmol mol^–1). After the fumigation, the net photosynthetic rate, stomatal conductance, transpiration rate, and F_v/F_m were reduced (p<0.05) for the three species. C. echinata was the most sensitive species and P. guajava cv. Paluma the most resistant.     

Changes Induced by Low Oxygen Concentration in Photosynthetic and Respiratory CO2 Exchange in Phosphate-Deficient Bean Leaves

Effect of phosphorus deficiency on photosynthetic and respiratory CO₂ exchanges were analysed in primary leaves of 2-week-old bean (Phaseolus vulgaris L. cv. Golden Saxa) plants under non-photorespiratory (2 % O₂) and photorespiratory (21 % O₂) conditions. Low P decreased maximum net photosynthetic rate (P_Nmax) and increased the time necessary to reach it. In the leaves of P-deficient plants the relative decrease of P_Nmax at 2 % O₂ was larger than at 21 % O₂. The results suggested the influence of photorespiration in the cellular turnover of phosphates.     

Different Responses of Norway Spruce Needles from Shaded and Exposed Crown Layers to the Prolonged Exposure to Elevated CO2 Studied by Various Chlorophyll a Fluorescence Techniques

The acclimation depression of capacity of photon utilisation in photochemical reactions of photosystem 2 (PS2) can develop already after three months of cultivation of the Norway spruces (Picea abies [L.] Karst.) under elevated concentrations of CO₂ (i.e., ambient, AC, + 350 µmol(CO₂) mol^–1 = EC) in glass domes with adjustable windows. To examine the role that duration of EC plays in acclimation response, we determined pigment contents, rate of photosynthesis, and parameters of chlorophyll a fluorescence for sun and shade needles after three seasons of EC exposure. We found responses of shaded and exposed needles to EC. Whereas the shaded needles still profited from the EC and revealed stimulated electron transport, for the exposed needles the stimulation of both electron transport activity and irradiance saturated rate of CO₂ assimilation (P _Nmax) under EC already disappeared. No signs of the PS2 impairment were observed as judged from high values of potential quantum yield of PS2 photochemistry (F_V/F_M) and uniform kinetics of Q_A reoxidation for all variants. Therefore, the long-term acclimation of the sun-exposed needles to EC is not necessarily accompanied with the damage to the PS2 reaction centres. The eco-physiological significance of the reported differentiation between the responses of shaded and sun exposed needles to prolonged EC may be in changed contribution of the upper and lower crown layers to the production activity of the tree. Whereas for the AC spruces, P _Nmax of shaded needles was only less than 25 % compared to exposed ones, for the EC spruces the P _Nmax of shaded needles reached nearly 40 % of that estimated for the exposed ones. Thus, the lower shaded part of the crown may become an effective consumer of CO₂.     

Different Mechanisms for Photosystem 2 Reversible Down-Regulation in Pumpkin and Soybean Leaves Under Saturating Irradiance

After saturating irradiation for 3 h (SI), the original fluorescence F₀ increased while the photosystem 2 (PS2) photochemical efficiency (F_v/F_m) declined significantly. These parameters could largely recover to the levels of dark-adapted leaves after 3 h of subsequent dark recovery. No net loss of the D1 proteins occurred after SI. Soybean and pumpkin leaves had different responses to SI. Low temperature fluorescence parameters, F₆₈₅ and F₆₈₅/F₇₃₅, decreased significantly in soybean leaves but not in pumpkin leaves. Part of the light-harvesting complex LHC2 dissociated from PS2 complexes in soybean leaves but not in pumpkin leaves, as shown by sucrose density gradient centrifugation and SDS-PAGE. The photon-saturated PS2 electron transport activity declined significantly in pumpkin thylakoids but not in soybean thylakoids. In addition, a large amount of phosphorylated D1 proteins was found in dark-adapted soybean leaves but not in dark-adapted pumpkin leaves. Hence at excessive irradiance soybean and pumpkin have the same protective strategy against photo-damage, reversible down-regulation of PS2, but two different mechanisms, namely the reversible down-regulation is related to the dissociation of LHC2 in soybean leaves but not in pumpkin leaves. This revised version was published online in August 2006 with corrections to the Cover Date.     

Characteristics of Photosynthetic Apparatus in Mn-Starved Maize Leaves

The effects of Mn-deficiency on CO₂ assimilation and excitation energy distribution were studied using Mn-starved maize leaves. Mn-deficiency caused about 70 % loss in the photon-saturated net photosynthetic rate (P _N) compared to control leaves. The loss of P _N was associated with a strong decrease in the activity of oxygen evolution complex (OEC) and the linear electron transport driven by photosystem 2 (PS2) in Mn-deficienct leaves. The photochemical quenching of PS2 (q_P) and the maximum efficiency of PS2 photochemistry (F_v/F_m) decreased significantly in Mn-starved leaves under high irradiance, implicating that serious photoinhibition took place. However, the high-energy fluorescence quenching (q_E) decreased, which was associated with xanthophyll cycle. The results showed that the pool of de-epoxidation components of the xanthophyll cycle was lowered markedly owing to Mn deficiency. Linear electron transport driven by PS2 de-creased significantly and was approximately 70 % lower in Mn-deficient leaves than that in control, indicating less trans-thylakoid pH gradient was built in Mn deficient leaves. We suggest that the decrease of non-radiative dissipation depending on xanthophyll cycle in Mn-starved leaves is a result of the deficiency of trans-thylakoid pH gradient.     

Responses of Tobacco Plantlets to Change of Irradiance During Transfer from in vitro to ex vitro Conditions

Chlorophyll a fluorescence kinetics, net photosynthetic rate (P _N), water relations, and photosynthetic pigment contents were studied during acclimation of in vitro grown tobacco to higher irradiance (HL; 700 mol m^–2 s^–1). Plantlets were grown on medium containing sucrose in glass vessels (G-plants) or in Magenta boxes (M-plants) with better CO₂ supply in the latter ones. The effect of HL was studied either (1) in plantlets grown under original in vitro conditions (closed vessels), (2) in in vitro plantlets exposed to ambient CO₂ concentration (covers removed), or (3) in plantlets transplanted to ex vitro into pots with sand and nutrient solution. Higher P _N, and fraction of closed photosystem 2 (PS2) centres (1 – q_P), and lower content of xanthophyll cycle pigments were found in M-plants compared to G-plants. HL treatment caused photoinhibition particularly in plants kept in closed vessels. This was indicated by the decrease in the ratio of F_v/F_m and by the increase in non-photochemical quenching, 1 – qp, and content of xanthophyll cycle pigments. Better CO₂ supply ensured by the removal of closure lead to the moderate reduction of symptoms of photoinhibition, although stomatal conductance (g _s), transpiration rate (E), and P _N were negatively affected. The main reason was the decrease in relative air humidity, which caused similar reduction of P _N, E, and g _s after the transfer of plantlets to ex vitro. Nevertheless, plant response to HL seemed not to be affected by any possible root injury caused by transfer to ex vitro. The differences in contents of xanthophyll cycle pigments, degree of de-epoxidation, P _N, and quenching parameters between M- and G-plantlets were still significant 7 d after ex vitro transfer and HL acclimation.     

Photosynthetic Characteristics of Non-Leaf Organs of Winter Wheat Cultivars Differing in Ear Type and their Relationship with Grain Mass Per Ear

Chlorophyll content, photosystem 2 functioning (F_v/F_m, F_v/F₀), activity of ribulose-1,5-bisphosphate carboxylase/oxygenase, and net photosynthetic rates (P _N) of flag leaf blade, sheath, peduncle, and ear organs were assessed in large-ear type (Pin 7) and small-ear type (ND93) wheat cultivars. Some differences were found in photosynthetic properties between different green plant parts, the values of all studied parameters in ear parts being higher in Pin7 than in ND93. Furthermore, ear surface areas and ear P _N in 26 wheat genotypes measured at anthesis showed highly significant positive correlation with grain mass per ear. Hence a greater capability of ear photosynthesis may result in a greater grain yield in large-ear type cultivars.     

Photosynthetic Performance of Ginkgo biloba L. Grown Under High and Low Irradiance

Diurnal variation in net photosynthetic rate (P _N) of three-year-old plants of Ginkgo biloba was studied under open, O (receiving full sunlight), net-shade, NS (40 % of photosynthetically active radiation, PAR), or greenhouse, G (25 % PAR) conditions. In all three conditions, P _N was higher in morning along with stomatal conductance (g _s), and intercellular CO₂ concentration (C _i), while leaf temperature and vapour pressure deficit were low. The O-plants exhibited a typical decline in P _N during midday, which was not observed in NS-plants. This indicated a possible photoinhibition in O-plants as the ratio of variable to maximum fluorescence (F_v/F_m) and photosystem 2 (PS2) yield (_PS2) values were higher in the NS- and G-plants. On the contrary, stomatal density and index, chlorophyll a/b ratio, leaf thickness, and density of mesophyll cells were greater in O-plants. Further, higher P _N throughout the day along with higher relative growth rate under NS as compared to O and G suggested the better efficiency of Ginkgo plants under NS conditions. Therefore, this plant species could be grown at 40 % irradiance to meet the ever-increasing demand of leaf and also to increase its export potential.     

强光和短期高浓度CO2对玉米和大豆光能转化效率的影响

盆栽和人工光源条件下,玉米叶片在普通空气中对强光照射不敏感,在高浓度CO2中强光照射0～5h后,光合速率(Pn)逐渐降低,无机磷(Pi)限制是其主要原因之一;大豆叶片在普通空气中受强光照射5h后,Fv／Fm、Pn、羧化效率(CE)和表观量子效率(AOY)明显降低,Fo升高,在高浓度CO2和强光下大豆Fo上升、Fv／Fm和气体交换参数下降的幅度减小。研究表明,高浓度CO2可减轻强光对植物尤其是C3植物光合功能的损伤,有限地缓解光抑制,但不能完全消除强光导致的大豆Pn和气孔导度(Gs)的降低。     

Role of Nitrate in Photosynthetic Electron Transport of Chlorella Vulgaris

Addition of nitrate to a suspension of NO₃ ^--depleted Chlorella vulgaris cells raised the O₂-evolving capacity of the organism by 60%. The rate of O₂-evolution under flash irradiation of the depleted cells was drastically reduced, which could be restored by addition of NO₃ ^-. The 2,5-dibromo-3-methyl-6-isopropyl-p-benzoquinone (DBMIB)-insensitive O₂-evolution, i.e., photosystem (PS) 2 activity of NO₃--depleted cells, showed a 75% stimulation by addition of NO₃ ^-. PS1-mediated electron transport was also stimulated (50%) by addition of NO₃ ^-. Fluorescence yields of the NO₃ ^--depleted cells were significantly reduced. A normal fluorescence response was restored by the addition of NO₃ ^-. The fluorescence yield of the NO₃ ^--depleted and DCMU-treated-cells increased significantly after addition of NO₃ ^- ions, indicating a further reduction of the primary acceptor of PS2 (Q). In addition, the low temperature fluorescence emission spectra showed that energy transfer to PS2 and PS1 was much higher when nitrate was present. Hence nitrate accelerates the light-induced charge transfer from the intact O₂-evolving system to the primary electron acceptor of PS2 and stimulates the PS1-mediated electron transport. This revised version was published online in June 2006 with corrections to the Cover Date.     

Diurnal and Seasonal Changes in Photosynthesis and Photosystem 2 Photochemical Efficiency in Prosopis juliflora Leaves Subjected to Natural Environmental Stress

The plants of Prosopis juliflora growing in northern India are exposed to large variations of temperature, vapour pressure deficits (VPD), and photosynthetic photon flux density (PPFD) throughout the year. Under these conditions P. juliflora had two short periods of leaf production, one after the winter season and second after summer, which resulted in two distinct even aged cohorts of leaves. In winter with cold nights (2–8 °C) and moderate temperatures during the day, the plants showed high rates of photosynthesis. In summer the midday temperatures often reached <45 °C and plants showed severe inhibition of photosynthesis. The leaves of second cohort appeared in July and showed typical midday depression of photosynthesis. An analysis of diurnal partitioning of the absorbed excitation energy into photochemistry showed that a smaller fraction of the energy was utilised for photochemistry and a greater fraction was dissipated thermally, further the photon utilisation for photochemistry and thermal dissipation is largely affected by the interaction of irradiance and temperature. The plants showed high photochemical efficiency of photosystem 2 (PS2) at predawn and very little photoinhibition in all seasons except in summer. The photoinhibition in summer was pronounced with very poor recovery during night. Since P. juliflora exhibited distinct pattern of senescence and production of new leaves after winter and summer stress period, it appeared that the ontogenic characteristic together with its ability for safe dissipation of excess radiant energy in P. juliflora contributes to its growth and survival.     

Developmental Change in CO2 Compensation Concentrations in Spartina alterniflora Results from Sigmoidal Photosynthetic CO2 Responses

We investigated seasonal patterns of photosynthetic responses to CO₂ concentrations in Spartina alterniflora Loisel, an aerenchymous halophyte grass, from a salt marsh of the Bay of Fundy (NB, Canada), and from plants grown from rhizome in controlled-environment chambers. From late May to August, CO₂ compensation concentrations () of field-grown leaves varied between 2.5–10.7 cm³(CO₂) m^–3, with a mean of 5.4 cm³(CO₂) m^–3. From September onwards field leaves showed CO₂ compensation concentrations from 6.6–21.1 cm³(CO₂) m^–3, with a mean of 13.1 cm³ m^–3 well into the C₃–C₄ intermediate range. The seasonal variability in did not result from changing respiration, but rather from a sigmoidal response of net photosynthetic rate (P _N) to applied CO₂ concentration, found in all tested leaves but which became more pronounced late in the season. One explanation for the sigmoidal response of P _N to external CO₂ concentration could be internal delivery of CO₂ from roots and rhizomes to bundle sheath cells via the aerenchyma, but the sigmoidal responses in S. alterniflora persisted out to the tips of leaves, while the aerenchyma extend only to mid-leaf. The sigmoidicity persisted when CO₂ response curves were measured from low to high CO₂, or from high to low CO₂, and even when prolonged acclimation times were used at each CO₂ concentration.     

Photosynthetic pigments and gas exchange during ex vitro acclimation of tobacco plants as affected by CO2 supply and abscisic acid

Nicotiana tabacum L. plantlets were grown in glass vessels or in Magenta boxes with better CO₂ supply. To improve the ex vitro transfer we tested application of abscisic acid and elevated CO₂ concentration. In the first two weeks after transfer, net photosynthetic rate, chlorophyll a+b content, and Chl a/b ratio were higher, and content of xanthophyll cycle pigments lower in M-plants than in G-plants, but during further growth the differences almost disappeared. ABA application alleviated the risk of wilting because it decreased stomatal conductance. The effect of ABA was enhanced under CE (28 days after transfer). In situ, P_N was always higher at CE than at CA, but when measured under CA, positive effect of CE was found 2 and 16 days after transfer in M-plants and only 16 days after transfer in G-plants. Slightly increased Chl a content was found in all ABA-treated plants, and in M-plants grown under CE. The content of xanthophyll cycle pigments was lower under CE compared to CA, and the lowest one was found in ABA-treated M-plants grown under CE. On the contrary, the degree of their deepoxidation (DEPS) was slightly higher in plants grown under CE. No significant effects of ABA-treatment or growth under CE on fluorescence kinetic parameters were found and inconsistent effects on photochemical activities. The photochemical efficiency of PS2 (variable to maximum fluorescence ratio, F_v/F_m) after ex vitro transfer was similar to that in in vitro grown plants. This together with the values of DEPS indicated that no photodamage during ex vitro transfer occurred. This revised version was published online in June 2006 with corrections to the Cover Date.     

Dissipation of Excess Energy in Mehler-Peroxidase Reaction in Rumex Leaves During Salt Shock

By measurement of gas exchange and chlorophyll fluorescence, the effects of salt shock on photosynthesis and the mechanisms to protect photosynthetic machinery against photodamage during salt shock were investigated in leaves of Rumex seedlings. Salt shock induced significant decrease in photosynthesis both in 21 and 2 % O₂. In 21 % O₂, quantum yield of photosystem 2 (PS2) electron transport (_PS2) decreased slightly and q_P remained constant, suggesting that the excitation pressure on PS2 did not increase during salt shock. In 2 % O₂, however, both _PS2 and q_P decreased significantly, suggesting that the excitation pressure on PS2 increased during salt shock. NPQ increased slightly in 21 % O₂ whereas it increased significantly in 2 % O₂. The data demonstrated that during salt shock a considerable electron flow was allocated to oxygen reduction in the Mehler-peroxidase reaction (MPR). Under high irradiance and in the presence of saturating CO₂, the susceptibility of PS2 to photoinhibition in salt-shocked leaves was increased when the electron flow to oxygen in MPR was inhibited in 2 % O₂. Hence, MPR is important in photoprotection of Rumex seedlings during salt shock.     

Factors Limiting Photosynthetic Recovery in Sweet Pepper Leaves After Short-Term Chilling Stress Under Low Irradiance

The effects of chilling treatment (4 °C) under low irradiance, LI (100 mol m^–2 s^–1) and in the dark on subsequent recovery of photosynthesis in chilling-sensitive sweet pepper leaves were investigated by comparing the ratio of quantum yields of photosystem (PS) 2 and CO₂ assimilation, _PS2/_CO2, measured in normal air (21 % O₂, NA) and low O₂-air (2% O₂, LOA), and by analyzing chlorophyll (Chl) a fluorescence parameters. Chilling treatment in the dark had little effect on F_v/F_m and _PS2/_CO2, but it caused the decrease of net photosynthetic rate (P _N) under saturating irradiance after 6-h chilling treatment, indicating that short-term chilling alone did not induce PS2 photoinhibition. Furthermore, photorespiration and Mehler reaction also did not obviously change during subsequent recovery after chilling stress in the dark. During chilling treatment under LI, there were obvious changes in F_v/F_m and _PS2/_CO2, determined in NA or LOA. F_v/F_m could recover fully in 4 h at 25 °C, and _PS2/_CO2 increased at the end of the treatment, as determined in both NA and LOA. During subsequent recovery, _PS2/_CO2 in LOA decreased faster than in NA. Thus the Mehler reaction might play an important role during chilling treatment under LI, and photorespiration was an important process during the subsequent recovery. The recovery of PN under saturating irradiance determined in NA and LOA took about 50 h, implying that there were some factors besides CO₂ assimilation limiting the recovery of photosynthesis. From the progress of reduced P700 and the increase of the Mehler reaction during chilling under LI we propose that active oxygen species were the factors inducing PS1 photoinhibition, which prevented the recovery of photosynthesis in optimal conditions because of the slow recovery of the oxidizable P700.     

Responses of Pea and Triticale Photosynthesis and Growth to Long-wave UV-B Radiation

Pea plants were more susceptible to long-wave UV-B irradiation (305 – 320 nm, 7.7 kJ m^–2 d^–1, 4 weeks) in comparison with the triticale. This difference was more apparent from the changes in total area of leaves and dry mass of shoots, rather than from the parameters of chlorophyll fluorescence and net photosynthetic rate.     

Chlorophyll Fluorescence and Photosynthetic Gas Exchange Responses to Irradiance of Tree of Heaven (Ailanthus Altissima) in Contrasting Urban Environments

Sun-and shade-adapted plants of Ailanthus altissima utilized thermal-dissipative photoprotection (NPQ) across a range of photosynthetic photon flux densities (PPFD), with higher NPQ and lower maximum quantum yield of photosystem 2 photochemistry (F_v/F_m) in sun-adapted individuals, suggesting increased engagement of antennae-based quenching. Photosynthetic quantum requirements (Q_req; number of photons per CO₂) were similar in sun and shade plants, but were low and comparable to forest understory species. Diurnal measurements showed that PPFDs in both habitats were consistently above photosynthetic compensation irradiance, and frequently exceeded saturating values. In addition, sun- and shade-adapted individuals possessed stomata that tracked short-term fluctuations in PPFD. Thus A. altissima may be unique in that it couples high, shade-plant like photosynthetic efficiency with high photosynthetic capacity in high-irradiance, while stomatal attributes that optimize water use efficiency are maintained in the shade. These features may contribute to success of A. altissima in establishing in disturbance-prone urban systems, and facilitate its spread into more PPFD-limited and competitive natural ecosystems.     

High Irradiance Induced Changes of Photosystem 2 in Young and Mature Needles of Cypress (Cupressus sempervirens L.)

Photoinhibition of photosynthesis was studied in young and mature detached sun needles of cypress under high irradiance (HI) of about 1 900 mol m^–2 s^–1. The degree of photoinhibition was determined by means of the ratio of variable to maximum chlorophyll (Chl) fluorescence (F_v/F_m) and electron transport measurements. Compared with the mature needles, the young needles, containing about half the amount of Chl a+b per unit area, exhibited a higher proportion of total carotenoids (Car) as xanthophyll cycle pigments and had an increased ratio of Car/Chl a+b. The potential efficiency of photosystem (PS) 2, F_v/F_m, markedly declined in HI-treated young needles without significant increase of F₀ level. In contrast, the F_v/F_m ratio declined with significant increase of F₀ level in mature needles. In isolated thylakoids, the rate of whole chain and PS2 activity markedly decreased in young HI-needles in comparison with mature needles. A smaller inhibition of PS1 activity was observed in both needles. In the subsequent dark incubation, fast recovery was found in both needle Types that reached maximum PS2 efficiencies similar to those observed in non-photoinhibited needles. The artificial exogenous electron donors DPC, NH₂OH, and Mn²⁺ failed to restore the HI-induced loss of PS2 activity in mature needles, while DPC and NH₂OH significantly restored it in young needles. Hence, HI-inactivation was on the donor side of PS2 in young needles and on the acceptor side of PS2 in mature needles. Quantification of the PS2 reaction centre proteins D1 and 33 kDa protein of water splitting complex following HI-exposure of needles showed pronounced differences between young and mature needles. The large loss of PS2 activity in HI-needles was due to the marked loss of D1 protein of the PS2 reaction centre in mature needles and of the 33 kDa protein in young needles.     

低剂量UV-B辐射对乌拉尔甘草叶片光合机构的影响

为了探讨植物叶片对UV-B辐射增强的响应机制,采用叶绿素荧光测定技术,分别测定在人工模拟低剂量UV-B(2.4μW/cm~2)辐射条件下乌拉尔甘草叶片的叶绿素荧光诱导动力曲线、初始荧光(F_0)、最大荧光(F_m)、光合机构比活性参数(ABS/RC、TR_o/RC和ET_o/RC)和性能指数等变化规律。结果表明:(1)低剂量UV-B辐射未引起甘草叶片O-J-I-P叶绿素荧光诱导曲线中的相数发生改变,UV-B辐射对PSⅡ的影响主要发生在其受体侧,而非供体侧;(2)低剂量UV-B辐射引起了甘草叶片光合系统F_v/F_m以及F_m、F_0的明显变化,同时也影响了光合机构的开放程度和电子从Q_A向Q_B传递效率,从而影响了光转化效率;相应性能指数(PI_(abs)和PI_(total))的改变亦验证了此结果。研究认为,低剂量UV-B辐射抑制乌拉尔甘草叶片光合系统Ⅱ受体侧Q_A至PQ之间的电子传递效率,从而影响了Q_A之后的光化学反应及非光化学反应。