Continuous recording of photochemical and non-photochemical chlorophyll fluorescence quenching with a new type of modulation fluorometer

A newly developed fluorescence measuring system is employed for the recording of chlorophyll fluorescence induction kinetics (Kautsky-effect) and for the continuous determination of the photochemical and non-photochemical components of fluorescence quenching. The measuring system, which is based on a pulse modulation principle, selectively monitors the fluorescence yield of a weak measuring beam and is not affected even by extremely high intensities of actinic light. By repetitive application of short light pulses of saturating intensity, the fluorescence yield at complete suppression of photochemical quenching is repetitively recorded, allowing the determination of continuous plots of photochemical quenching and non-photochemical quenching. Such plots are compared with the time courses of variable fluorescence at different intensities of actinic illumination. The differences between the observed kinetics are discussed. It is shown that the modulation fluorometer, in combination with the application of saturating light pulses, provides essential information beyond that obtained with conventional chlorophyll fluorometers.     

Detection of rapid induction kinetics with a new type of high-frequency modulated chlorophyll fluorometer

A newly developed modulation fluorometer is described which operates with 1 sec light pulses from a light-emitting diode (LED) at 100 KHz. Special amplification circuits assure a highly selective recording of pulse fluorescence signals against a vast background of non-modulated light. The system tolerates ratios of up to 1:10⁷ between measuring light and actinic light. Thus it is possible to measure the dark fluorescence yield and record the kinetics of light-induced changes. A high time resolution allows the recording of the rapid relaxation kinetic following a saturating single turnover flash. Examples of system performance are given. It is shown that following a flash the reoxidation kinetics of photosystem II acceptors are slowed down not only by the inhibitor DCMU, but by a number of other treatments as well. From a light intensity dependency of the induction kinetics the existence of two saturated intermediate levels (I₁ and I₂) is apparent, which indicates the removal of three distinct types of fluorescence quenching in the overall fluorescence rise from F₀ to F_max.Abbreviations Q_A and Q_B consecutive electron acceptors of photosystem II - PS II photosystem II - P 680 reaction center chlorophyll of photosystem II - F₀ minimum fluorescence yield following dark adaptation - F_max maximum fluorescence yield - DCMU 3-(3, 4-dichlorophenyl)-1, 1-dimethyl-urea - DCCD N,N-dicyclohexylcarbodiimide - PQ plastoquinone - DAD diaminodurene Dedicated to Prof. L.N.M. Duysens on the occasion of his retirement.     

A solid-state,portable instrument for measurement of chlorophyll luminescence induction in plants

A newly developed compact instrument is described for the measurement of chlorophyll luminescence induction in plants. The instrument operates with a pulsed light emitting diode (LED) as light source and a photodiode as luminescence detector. A special emitter-detector geometry provides for high irradiance of the sample and efficient collection of luminescence by the detector. With insertion of appropriate filters the same probe is also suited for measuring prompt chlorophyll fluorescence. The instrument shows considerable flexibility with respect to pulse frequency, relative lengths of light/dark intervals and luminescence sampling periods. Due to a selective amplifier system only that part of luminescence is processed which is induced by the individual excitation pulses. By this approach, the problem of slow phase accumulation, encountered with conventional phosphoroscopes, is eliminated. Some examples are given for system operation, demonstrating satisfactory performance in measurements with intact leaves and isolated chloroplasts.     

Sensitivity of the relative Fpl level of chlorophyll fluorescence induction in leaves to the heat stress

The (F_pl-F_o)/F_v value of the fluorescence induction curve is shown to be a more suitable parameter to detect a wider range of heat stress damage to thylakoid membranes as compared to quantities t _1/2 (time of fluorescence rise from F_o to (F_o+F_m)/2 level) and % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0Jf9crFfpeea0xh9v8qiW7rqqrFfpeea0xe9Lq-Jc9% vqaqpepm0xbba9pwe9Q8fs0-yqaqpepae9pg0FirpepeKkFr0xfr-x% fr-xb9adbaqaaeGaciGaaiaabeqaamaabaabaaGcbaWaa0aaaeaacq% aHepaDaaaaaa!39D5!\[\overline \tau \] (the fluorescence induction time defined as the area above the induction curve normalized to F_v=1). A method for exact and automatic F_pl determination is presented.A break point in the quality and behaviour of the fluorescence induction curve of barley leaves incubated at 49°C was reached at the moment (about 240 s) when the transformation of PS II active (Q_B-reducing) to PS II inactive (Q_B-non-reducing) centres was completed. The meaning of the standard F_v and F_v/F_m parameter was then changed.The method of F_pl determination described here may help to increase the analytical value of the standard chlorophyll fluorometers.Abbreviations F_o initial fluorescence - F_m maximal fluorescence - F_pl fluorescence at first inflection point (plateau) - F_v variable fluorescence (F_v=F_m–F_o) - PSM plant stress meter - SD standard deviation     

The fast and slow kinetics of chlorophyll a fluorescence induction in plants, algae and cyanobacteria: a viewpoint

The light-induced/dark-reversible changes in the chlorophyll (Chl) a fluorescence of photosynthetic cells and membranes in the μs-to-several min time window (fluorescence induction, FI; or Kautsky transient) reflect quantum yield changes (quenching/de-quenching) as well as changes in the number of Chls a in photosystem II (PS II; state transitions). Both relate to excitation trapping in PS II and the ensuing photosynthetic electron transport (PSET), and to secondary PSET effects, such as ion translocation across thylakoid membranes and filling or depletion of post-PS II and post-PS I pools of metabolites. In addition, high actinic light doses may depress Chl a fluorescence irreversibly (photoinhibitory lowering; q(I)). FI has been studied quite extensively in plants an algae (less so in cyanobacteria) as it affords a low resolution panoramic view of the photosynthesis process. Total FI comprises two transients, a fast initial (OPS; for Origin, Peak, Steady state) and a second slower transient (SMT; for Steady state, Maximum, Terminal state), whose details are characteristically different in eukaryotic (plants and algae) and prokaryotic (cyanobacteria) oxygenic photosynthetic organisms. In the former, maximal fluorescence output occurs at peak P, with peak M lying much lower or being absent, in which case the PSMT phases are replaced by a monotonous PT fluorescence decay. In contrast, in phycobilisome (PBS)-containing cyanobacteria maximal fluorescence occurs at M which lies much higher than peak P. It will be argued that this difference is caused by a fluorescence lowering trend (state 1 → 2 transition) that dominates the FI pattern of plants and algae, and correspondingly by a fluorescence increasing trend (state 2 → 1 transition) that dominates the FI of PBS-containing cyanobacteria. Characteristically, however, the FI pattern of the PBS-minus cyanobacterium Acaryochloris marina resembles the FI patterns of algae and plants and not of the PBS-containing cyanobacteria.     

Evaluation of a technique for the measurement of chlorophyll fluorescence from leaves exposed to continuous white light

Abstract An instrument for the generation and measurement of modulated chlorophyll fluorescence signals from leaves exposed to continuous, highintensity white light is described. Modulated fluorescence is generated in the leaf by pulsed diodes emitting low-intensity yellow radiation and is detected with a photodiode whose output is fed to an amplifier locked in to the frequency of the lightemitting diodes. Comparisons are made between the modulated fluorescence signals measured with this instrument and the continuous fluorescence signals emitted from dark-adapted leaf tissue and isolated thylakoids when photosynthetic activity is induced by exposure to a range of intensities of continuous broad-band, blue-green light. The modulated fluorescence signals were similar to the continuous fluorescence signals, but they were not always identical. The small differences between the two signals are mainly attributable to differences in the populations of chloroplasts being monitored in the two measurements as a result of differential penetration of the modulated and actinic light sources into the sample.     

PAM fluorometer based on medium-frequency pulsed Xe-flash measuring light: A highly sensitive new tool in basic and applied photosynthesis research

A newly developed modulation fluorometer is described which employs repetitive 1 s Xe-flashes for excitation light. Similar to the standard PAM Chlorophyll Fluorometer, which uses 1 s LED pulses for measuring light, the integrated measuring light intensity is sufficiently low to monitor the dark-fluorescence level, F_o. The maximal fluorescence yield, F_m, can be determined with high selectivity upon application of a saturating light pulse. The Xe-PAM displays exceptionally high sensitivity, enabling quenching analysis at chlorophyll concentrations as low as 1 g/l, thus allowing to assess photosynthesis of phytoplankton in natural waters like lakes, rivers and oceans. Due to high flexibility in the choice of excitation and emission wavelengths, this system also provides the experimental basis for a thorough study of fluorescence and photosynthesis properties of various algae classes with differing antenna organisation. By appropriate modifications, the instrument may as well be used to measure with great sensitivity and selectivity other types of fluorescence (e.g. NADPH-fluorescence), as well as light-scattering and absorbance changes.     

盐胁迫对植物叶绿素荧光影响的研究进展

盐胁迫是制约植物生长发育的主要非生物胁迫之一, 研究植物的耐盐机理对开发和有效利用盐碱地有重要的意义。叶绿素荧光动力技术作为研究植物光合生理状况及植物与逆境胁迫关系的理想方法, 可表明外界胁迫环境对植物光合器官的伤害程度。通过总结性阐述盐胁迫对植物叶绿素荧光的影响, 分别从盐分类型、植物类型、光照强度以及盐旱交互作用等方面分析了植物叶绿素荧光对盐胁迫的响应, 进而反映盐胁迫对植物光合能力的影响程度, 并提出增强植物抗盐性的途径, 包括施加外源物质、利用转基因技术、真菌的协同效应和培育耐盐品种。最后对叶绿素荧光动力技术在抗盐胁迫的运用前景进行了展望, 提出了当前研究需要解决的问题, 旨在为提高植物耐盐能力提供一定的理论依据。     

Excitation kinetics during induction of chlorophyll <Emphasis Type="Italic">a</Emphasis> fluorescence

We present a chlorophyll fluorometer module system which adapts the intensity to the individual leaf sample by adjusting the quantum flux density of the excitation light so that the fluorescence signal is kept constant. This is achieved by means of a feedback power adjustment of the fluorescence exciting laser diode. Thus, the intensity of the excitation light is adapted to the actual need of a particular sample for quantum conversion without applying exaggeratedly high quantum flux density. We demonstrate the influence of the initial laser power chosen at the onset of irradiation and kept constant during fluorescence rise transient within the first second. Examples are shown for measuring upper and lower leaf sides, a single leaf with different pre-darkening periods, as well as yellow, light green and dark green leaves. The novel excitation kinetics during the induction of chlorophyll fluorescence can be used to study the yield and regulation of photosynthesis and its related non-photochemical processes for an individual leaf. It allows not only to sense the present state of pre-darkening or pre-irradiation but also the light environment the leaf has experienced during its growth and development. Thus, the individual physiological capacity and plasticity of each leaf sample can be sensed being of high importance for basic and applied ecophysiological research which makes this new methodology both innovative and informative.     

Effects of solar UV stress on chlorophyll fluorescence kinetics of intertidal macroalgae from southern Spain: a case study in Gelidium species

Photoinhibition and recovery kinetics after short exposure to solar radiation following three different irradiance treatments of irradiances (PAR, PAR+UVA and PAR+UVA+UVB) was assessed in two intertidal species of the genus Gelidium, Gelidium sesquipedale and G. latifolium, collected from Tarifa (southern Spain) using in vivo chlorophyll fluorescence (PAM fluorometry). After 3 h UV radiation exposure, optimal quantum efficiency (Fv/Fm) in G. sesquipedale decreased between 25 and 35% relative to the control. Under PAR alone, values decreased to 60%. In G. latifolium, photoinhibition did not exceed 40%. Similar results were found for the effective quantum yield (ΔF/Fm′), however, no marked differences in relation to light treatments were seen. When plants were shaded for recovery from stress, only in G. latifolium a significant increase in photosynthesis was observed (between 80 and 100% of control). In contrast, photosynthesis of G. sesquipedale suffered a chronic photoinhibition or photodamage under the three light irradiances. Full solar radiation (PAR+UVA+UVB) affected also the electron transport rate in both species. Here, initial slopes of electron transport vs. irradiance curves decreased up to 60% of controls. Although the recovery kinetic under PAR+UVA+UVB conditions was delayed in G. latifolium, after 24 h recovery this species reached significantly higher than G. sesquipedale. PAR impaired electron trasport only in G. sesquipedale. Overall, both species are characterized by different capacity to tolerate enhanced solar radiation. G. latifolium is a sun adapted plant, well suited to intertidal light conditions, whereas G. sesquipedale, growing at shaded sites in the intertidal zone, is more vulnerable to enhanced UV radiation. This revised version was published online in June 2006 with corrections to the Cover Date.     

Effects of frost hardening, dehardening and freezing stress on in vivo chlorophyll fluorescence of seedlings of Scots pine (Pinus sylvestris L.)

Abstract. The kinetics of in vivo chlorophyll fluorescence of photosystem II (PS II) was measured at room temperature and 77 K during frost hardening of seedlings of Scots pine (Pinus sylvestris L.), and after exposure of frost-hardened shoots to sub-freezing temperatures. A more pronounced decrease in variable fluorescence yield for the upper exposed than for the lower shaded surface of the needles suggested that some photoinhibition occurred during prolonged frost hardening at 50 μmol photons m^?2 s^?1 and 4°C. Reversible inhibition of photosynthesis after exposure to sub-freezing temperatures was initially manifested as an increase of steady-state energy-dependent fluorescence quenching (q_E) and a reduction in the rate of O₂ evolution. Further inhibition after treatment at still lower temperatures caused a progressive decline of steady-state photochemical quenching (q_Q) and the rate of O₂ evolution, whereas q_E remained high. This implies an inactivation of enzymes in the photosynthetic carbon reduction cycle decreasing the consumption of ATP and NADPH, which is likely to cause an increase of membrane energization and a reduction of the primary electron acceptor (Q_A) of PS II. Alternatively, the changes in q_Q and q_E might be attributed to an inhibition of photophosphorylation. Severe, irreversible damage to photosynthesis resulted in a suppression of q_E and of variable fluorescence yield, probably because the photochemical efficiency of PS II was impaired. Changes in the fast fluorescence kinetics at room temperature after severe freezing damage were interpreted as an inhibition of the electron flow from Q_A to the plastoquinone pool. It is suggested that irreversible freezing injury to needles of frost-hardened P. sylvestris causes damage to the Q_B,-protein.     

Evaluation of instant light‐response curves of chlorophyll fluorescence parameters obtained with a portable chlorophyll fluorometer on site in the field

Miniaturized pulse‐amplitude modulated photosynthesis yield analysers are primarily designed for measuring effective quantum yield (ΔF/F_m′) of photosystem II under momentary ambient light conditions in the field. Although this provides important ecophysiological information, it is often necessary to learn more about the potential intrinsic capacities of leaves by measuring light‐response curves. Thus, instruments provide light‐curve programmes, where light intensities are increased in short intervals and instant light‐response curves are recorded within a few minutes. This method can be criticized because photosynthesis will most likely not be in steady state. This technical report shows that with the appropriate precautions instant light curves can nevertheless provide reliable information about cardinal points of photosynthesis. First, the geometry of the light source of the instrument in relation to the quantum sensor must be considered and quantum sensor readings must be corrected. Second, the measurements of the light‐response curves must be compared with readings of effective quantum yield of photosystem II under ambient light conditions where photosynthesis is in steady state. This may show that in the critical range of the light curves either both measurements perfectly coincide or are offset against each other by a constant value (examples are given here). In the first case results of light curves can be taken at face values, and in the second case a simple correction can be applied. With these precautions and careful interpretations instant light‐response curves can be an enormous advantage in ecophysiological field work.     

Resolution of components of non-photochemical chlorophyll fluorescence quenching in barley leaves

Non-photochemical chlorophyll fluorescence quenching (qN) in barley leaves has been analysed by monitoring its relaxation in the dark, by applying saturating pulses of light. At least three kinetically distinct phases to qN recovery are observed, which have previously been identified (Quick and Stitt 1989) as being due to high-energy state quenching (fast), excitation energy redistribution due to a state transition (medium) and photoinhibition (slow). However, measurements of chlorophyll fluorescence at 77 K from leaf extracts show that state transitions only occur in low light conditions, whereas the medium component of qN is very large in high light. The source of that part of the medium component not accounted for by a state transition is discussed.Abbreviations ATP adenosine 5-triphosphate - DCMU 3[3,4-dichlorophenyl]-1,1 dimethylurea - pH trans-thylakoid pH gradient - F_o, F_m room-temperature chlorophyll fluorescence yield with all reaction centres open, closed - F_v variable fluorescence = F_m–F_o - LHC II Light harvesting complex II - PS I, PS II Photosystem I, II - P700, P680 primary donor in photosystem I, II - qP photochemical quenching of variable fluorescence - qN non-photochemical quenching of variable fluorescence - qN_e, qN_t, qN_i non-photochemical quenching due to high energy state, state transition, photoinhibition - qN_f, qN_m, qN_s components of qN relaxing fast, medium, slow - q_r quenching of r relative to the dark state - tricine N-tris[hydroxymethyl]methylglycine - r ratio of fluorescence maximum from photosystem II to that from photosystem I at 77 K     

Light-induced fluorescence quenching in the light-harvesting chlorophyll a/b protein complex

Summary Irradiation of the principal photosystem II light-harvesting chlorophyll-protein antenna complex, LHC II, with high light intensities brings about a pronounced quenching of the chlorophyll fluorescence. Illumination of isolated thylakoids with high light intensities generates the formation of quenching centres within LHC II in vivo, as demonstrated by fluorescence excitation spectroscopy. In the isolated complex it is demonstrated that the light-induced fluorescence quenching: a) shows a partial, biphasic reversibility in the dark; b) is approximately proportional to the light intensity; c) is almost independent of temperature in the range 0–30°C; d) is substantially insensitive to protein modifying reagents and treatments; e) occurs in the absence of oxygen. A possible physiological importance of the phenomenon is discussed in terms of a mechanism capable of dissipating excess excitation energy within the photosystem II antenna.Abbreviations chla chlorophyll a - chlb chlorophyll b - F₀ fluorescence yield with reaction centers open - F_m fluorescence yield with reaction centres closed - F_i fluorescence at the plateau level of the fast induction phase - LHC II light-harvesting chlorophyll a/b protein complex II - PS II photosystem II - PSI photosystem I - Tricine N-[2-hydroxy-1,1-bis(hydroxymethyl)ethyl]glycine     

高温胁迫对新疆榛光合参数和叶绿素荧光特性的影响

在5个温度梯度处理下,研究高温胁迫对4种新疆榛光合参数和叶绿素荧光特性的影响.结果表明:随着温度从25℃持续升高至45℃,新疆榛叶片的净光合速率、气孔导度、胞间CO2浓度、水分利用效率和光能利用效率逐渐降低,且在35～ 45℃之间降幅最大;光系统Ⅱ的实际光化学效率、电子传递速率和光化学猝灭系数随温度的升高缓慢上升,至35℃后急速下降;蒸腾耗水和热耗散随温度的升高而增大.4种新疆榛品种中,新榛3号的光合作用对高温的耐受力较高,属耐热性品种.     

pH dependent chlorophyll fluorescence quenching in spinach thylakoids from light treated or dark adapted leaves

The pH dependence of maximum chlorophyll fluorescence yield (F_m) was examined in spinach thylakoids in the presence of nigericin to dissipate the transthylakoid pH gradient. 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU) was present to eliminate photochemical quenching. Thylakoids were prepared from dark adapted leaves (dark thylakoids) or preilluminated leaves (light thylakoids). In the latter there had been approximately 50% conversion of the xanthophyll violaxanthin to zeaxanthin, while no conversion had occurred in the former. In the presence of a reductant such as ascorbate, antimycin A sensitive quenching was observed (half maximal quenching at 5 M), whose pH dependence differed between the two types of thylakoid. Preillumination of leaves resulted in more quenching at pH values where very little quenching was observed in dark thylakoids (pH 5–7.6). This was similar to activation of high-energy-state quenching (qE) observed previously (Rees D, Young A, Noctor G, Britton G and Horton P (1989) FEBS Lett 256: 85–90). Thylakoids isolated from preilluminated DTT treated leaves, that contained no zeaxanthin, behaved like dark thylakoids. A second form of quenching was observed in the presence of ferricyanide, that could be reversed by the addition of ascorbate. This was not antimycin A sensitive and showed the same pH dependence in both types of thylakoid. The former type of quenching, but not the latter, showed similar low temperature fluorescence emission spectra to qE, and was considered to occur by the same mechanism.Abbreviations DCMU 3(3,4-dichlorophenyl)-1,1-dimethylurea - DTT dithiothreitol - EDTA Ethylenediaminetetra-acetic acid - F₀ dark level fluorescence yield - F_m maximum fluorescence yield - F_v/F_m ratio of variable to total fluorescence yield - Hepes 4-(2-hydroxyethyl)1-piperazineethanesul-phonic acid - Mes 2-(N-morpholino) ethanesulfonate - pH transthylakoid pH gradient - PS I Photosystem I - PS II Photosystem II - Q_A primary stable electron acceptor of Photosystem II - qE high-energy-state fluorescence quenching     

Effect of salt stress on photosynthesis and chlorophyll fluorescence in Medicago truncatula

In the present study, photosynthetic parameters including gas exchanges, pigment contents, and chlorophyll fluorescence, were compared in two contrasting local Medicago truncatula lines TN6.18 and TN8.20, in response to salt added to the nutrient solution. Plants were cultivated under symbiotic nitrogen fixation (SNF) after inoculation with a reference strain Sinorhizobium meliloti 2011, a very tolerant strain to salinity (700 mM NaCl), and grown in a controlled glasshouse. On one month old plants (with active SNF), salt treatment (75 mM NaCl) was gradually applied. Photosynthesis, assimilating pigments and chlorophyll fluorescence were monitored throughout the experiment during both short and long terms, compared to control (non-saline) conditions. A genotypic variation in salt tolerance was found; TN6.18 was the more sensitive to salinity. The relative tolerance of TN8.20 was concomitant with the highest photochemical quenching coefficient (q_P) affecting the maximum quantum yield of PSII (Y); the real quantum yield (?_exc) was the most affected in the sensitive line. Moreover, stomatal and PSII reaction centers activities differed clearly between the studied lines. We found that the effect of salinity on photosynthesis of M. truncatula was related to PSII activity reduction rather than to stomatal conductance limitation. Photosynthesis was reduced by the inhibition of CO₂ assimilation caused by PSII damage. This was clearly estimated by the Y, ?_exc and especially by the quantum yield of electron transport of PSII (Φ_PSII). Thus, on the basis of our results on the two local M. truncatula lines, we recommend the use of chlorophyll fluorescence as non-destructive screening method to discriminate susceptible and resistant legumes to salt stress.     

Relaxation of the non-photochemical chlorophyll fluorescence quenching in diatoms: kinetics,components and mechanisms

Diatoms are especially important microorganisms because they constitute the larger group of microalgae. To survive the constant variations of the light environment, diatoms have developed mechanisms aiming at the dissipation of excess energy, such as the xanthophyll cycle and the non-photochemical chlorophyll (Chl) fluorescence quenching. This contribution is dedicated to the relaxation of the latter process when the adverse conditions cease. An original nonlinear regression analysis of the relaxation of non-photochemical Chl fluorescence quenching, qN, in diatoms is presented. It was used to obtain experimental evidence for the existence of three time-resolved components in the diatom Phaeodactylum tricornutum: qNf, qNi and qNs. qNf (s time-scale) and qNs (h time-scale) are exponential in shape. By contrast, qNi (min time-scale) is of sigmoidal nature and is dominant among the three components. The application of metabolic inhibitors (dithiothreitol, ammonium chloride, cadmium and diphenyleneiodonium chloride) allowed the identification of the mechanisms on which each component mostly relies. qNi is linked to the relaxation of the ΔpH gradient and the reversal of the xanthophyll cycle. qNs quantifies the stage of photoinhibition caused by the high light exposure, qNf seems to reflect fast conformational changes within thylakoid membranes in the vicinity of the photosystem II complexes.     

Drought and chlorophyll fluorescence in field-grown potato (Solanum tuberosum)

Light interception, stomatal conductance and chlorophyll fluorescence were measured in potato ( Solanum tuberosum L.) grown either irrigated, or droughted from the time of plant emergence. Compared with the irrigated treatment, drought reduced both light interception and stomatal conductance. In both treatments, the yields of variable fluorescence in the dark- and light-adapted states (F_y/F_m and F'_v/F'_m, respectively) were negatively correlated with photosynthetic photon flux density (PPFD) and mirrored daytime changes in PPFD. Photochemical quenching was positively correlated with PPFD, but the dominant effect of F'_v/F'_m resulted in a decrease in the quantum yield of photosystem II (PSII) electron transport with increasing PPFD.
Drought had no significant effect on the functioning of PSII and the balance between photochemical and non-photochemical quenching was unaffected. Non-photochemical quenching was not increased by drought and the quantum yield of PSII electron transport was unaffected. It is concluded that, in leaves of droughted plants, excess energy, resultant of stomatal limitation of photosynthesis, was dissipated by photochemical quenching such as increased photorespiration.