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Journal ArticleDOI

Chlorophyll Fluorescence Parameters: The Definitions, Photosynthetic Meaning, and Mutual Relationships

01 Mar 2002-Photosynthetica (Institute of Experimental Botany)-Vol. 40, Iss: 1, pp 13-29
TL;DR: In this article, a review of Chlorophyll fluorescence parameters derived from the slow (long-term) induction kinetics of modulated Chl a fluorescence are reviewed and analyzed with respect to their application in photosynthesis research.
Abstract: Chlorophyll fluorescence parameters (Chl FPs) derived from the slow (long-term) induction kinetics of modulated Chl a fluorescence are reviewed and analysed with respect to their application in photosynthesis research. Only four mutually independent Chl FPs, calculated from values of five essential Chl fluorescence (ChlF) yields, are distinguished as the basic ones. These are: the maximum quantum yield of PS2 photochemistry (ΦPO), the photochemical quenching of variable ChlF (qP), the non-photochemical quenching of variable ChlF (qN), and the relative change of minimum ChlF (qO). ΦPO refers to the dark-adapted state of a thylakoid membrane, qP, qN and qO characterise the light-adapted state. It is demonstrated that all other Chl FPs can be determined using this quartet of parameters. Moreover, three FPs related to the non-radiative energy dissipation within thylakoid membranes are evaluated, namely: the non-photochemical ChlF quenching (NPQ), the complete non-photochemical quenching of ChlF (qCN), and the effective quantum yield of non-photochemical processes in PS2 (ΦN). New FPs, the total quenching of variable ChlF (qTV) and the absolute quenching of ChlF (qA) which allow to quantify co-action of the photochemical and non-photochemical processes during a light period are defined and analysed. The interpretation of Chl FPs and recommendations for their application in the photosynthesis research are also given. Some alternative FPs used in the laboratory practice have only an approximate character and can lead to incorrect conclusions if applied to stressed plants. They are reviewed and compared with the standard ones. All formulae and conclusions discussed herein are verified using experimental values obtained on young seedlings of the Norway spruce (Picea abies [L.] Karst.).
Citations
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Journal ArticleDOI
TL;DR: Progress made during the last two decades in producing transgenic lines of different C3 crops with enhanced photosynthetic performance is discussed, which was reached by either the overexpression of C3 enzymes or transcription factors or the incorporation of genes encoding C4 enzymes into C3 plants.
Abstract: Stressful environments such as salinity, drought, and high temperature (heat) cause alterations in a wide range of physiological, biochemical, and molecular processes in plants. Photosynthesis, the most fundamental and intricate physiological process in all green plants, is also severely affected in all its phases by such stresses. Since the mechanism of photosynthesis involves various components, including photosynthetic pigments and photosystems, the electron transport system, and CO2 reduction pathways, any damage at any level caused by a stress may reduce the overall photosynthetic capacity of a green plant. Details of the stress-induced damage and adverse effects on different types of pigments, photosystems, components of electron transport system, alterations in the activities of enzymes involved in the mechanism of photosynthesis, and changes in various gas exchange characteristics, particularly of agricultural plants, are considered in this review. In addition, we discussed also progress made during the last two decades in producing transgenic lines of different C3 crops with enhanced photosynthetic performance, which was reached by either the overexpression of C3 enzymes or transcription factors or the incorporation of genes encoding C4 enzymes into C3 plants. We also discussed critically a current, worldwide effort to identify signaling components, such as transcription factors and protein kinases, particularly mitogen-activated protein kinases (MAPKs) involved in stress adaptation in agricultural plants.

1,435 citations


Cites background from "Chlorophyll Fluorescence Parameters..."

  • ...Additional key words: drought; fluorescence; gas exchange; heat; photosynthesis; photosynthetic pigments; salinity, salinity stress....

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  • ...It gives a valuable insight into exploitation of the excitation energy by PSII, and indirectly by the other protein complexes of the thylakoid membranes (Walker 1987, Roháček 2002), particularly in plants exposed to stressful conditions....

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Journal ArticleDOI
TL;DR: The basis of photosynthetic acclimation and its optical signals is presented, the physical and physiological basis of ChlF is introduced from the molecular to the leaf level and beyond, and PAM and SIF methodology are introduced.
Abstract: subcellular levels. ChlF is now measurable from remote sensing platforms. This provides a new optical means to track photosynthesis and gross primary productivity of terrestrial ecosystems. Importantly, the spatiotemporal and methodological context of the new applications is dramatically different compared with most of the available ChlF literature, which raises a number of important considerations. Although we have a good mechanistic understanding of the processes that control the ChlF signal over the short term, the seasonal link between ChlF and photosynthesis remains obscure. Additionally, while the current understanding of in vivo ChlF is based on pulse amplitude-modulated (PAM) measurements, remote sensing applications are based on the measurement of the passive solar-induced chlorophyll fluorescence (SIF), which entails important differences and new challenges that remain to be solved. In this review we introduce and revisit the physical, physiological, and methodological factors that control the leaf-level ChlF signal in the context of the new remote sensing applications. Specifically, we present the basis of photosynthetic acclimation and its optical signals, we introduce the physical and physiological basis of ChlF from the molecular to the leaf level and beyond, and we introduce and compare PAM and SIF methodology. Finally, we evaluate and identify the challenges that still remain to be answered in order to consolidate our mechanistic understanding of the remotely sensed SIF signal.

714 citations


Cites background from "Chlorophyll Fluorescence Parameters..."

  • ...Further photochemical and non-photochemical quenching parameters and process quantum yields can be found elsewhere (e.g. Rohácek, 2002; Krause and Jahns, 2004; Schreiber, 2004; Baker, 2008; PorcarCastell, 2011)....

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Journal ArticleDOI
TL;DR: A role for activase in limiting photosynthesis at high temperature is supported, as the temperature response of Pn, Rubisco activation, chlorophyll fluorescence, and the activities of Rubisco and Rubisco activase were examined in species from contrasting environments.
Abstract: Inhibition of net photosynthesis (Pn) by moderate heat stress has been attributed to an inability of Rubisco activase to maintain Rubisco in an active form. To examine this proposal, the temperature response of Pn, Rubisco activation, chlorophyll fluorescence, and the activities of Rubisco and Rubisco activase were examined in species from contrasting environments. The temperature optimum of Rubisco activation was 10°C higher in the creosote bush (Larrea tridentata) compared with the Antarctic hairgrass (Deschampsia antarctica), resembling the temperature response of Pn. Pn increased markedly with increasing internal CO2 concentration in Antarctic hairgrass and creosote bush plants subjected to moderate heat stress even under nonphotorespiratory conditions. Nonphotochemical quenching of chlorophyll fluorescence, the effective quantum yield of photochemical energy conversion (ΔF/Fm′) and the maximum yield of PSII (Fv/Fm) were more sensitive to temperature in Antarctic hairgrass and two other species endemic to cold regions (i.e. Lysipomia pumila and spinach [Spinacea oleracea]) compared with creosote bush and three species (i.e. jojoba [Simmondsia chinensis], tobacco [Nicotiana tabacum], and cotton [Gossypium hirsutum]) from warm regions. The temperature response of activity and the rate of catalytic inactivation of Rubisco from creosote bush and Antarctic hairgrass were similar, whereas the optimum for ATP hydrolysis and Rubisco activation by recombinant creosote bush, cotton, and tobacco activase was 8°C to 10°C higher than for Antarctic hairgrass and spinach activase. These results support a role for activase in limiting photosynthesis at high temperature.

450 citations

Journal ArticleDOI
TL;DR: In this article, a practical guide to the measurement of the different chlorophyll (Chl) fluorescence parameters and gives examples of their development under high-irradiance stress.
Abstract: This contribution is a practical guide to the measurement of the different chlorophyll (Chl) fluorescence parameters and gives examples of their development under high-irradiance stress. From the Chl fluorescence induction kinetics upon irradiation of dark-adapted leaves, measured with the PAM fluorometer, various Chl fluorescence parameters, ratios, and quenching coefficients can be determined, which provide information on the functionality of the photosystem 2 (PS2) and the photosynthetic apparatus. These are the parameters Fv, Fm, F0, Fm′, Fv′, NF, and ΔF, the Chl fluorescence ratios Fv/Fm, Fv/F0, ΔF/Fm′, as well as the photochemical (qP) and non-photochemical quenching coefficients (qN, qCN, and NPQ). qN consists of three components (qN = qE + qT + qI), the contribution of which can be determined via Chl fluorescence relaxation kinetics measured in the dark period after the induction kinetics. The above Chl fluorescence parameters and ratios, many of which are measured in the dark-adapted state of leaves, primarily provide information on the functionality of PS2. In fully developed green and dark-green leaves these Chl fluorescence parameters, measured at the upper adaxial leaf side, only reflect the Chl fluorescence of a small portion of the leaf chloroplasts of the green palisade parenchyma cells at the upper outer leaf half. Thus, PAM fluorometer measurements have to be performed at both leaf sides to obtain information on all chloroplasts of the whole leaf. Combined high irradiance (HI) and heat stress, applied at the upper leaf side, strongly reduced the quantum yield of the photochemical energy conversion at the upper leaf half to nearly zero, whereas the Chl fluorescence signals measured at the lower leaf side were not or only little affected. During this HL-stress treatment, qN, qCN, and NPQ increased in both leaf sides, but to a much higher extent at the lower compared to the upper leaf side. qN was the best indicator for non-photochemical quenching even during a stronger HL-stress, whereas qCN and NPQ decreased with progressive stress even though non-photochemical quenching still continued. It is strongly recommended to determine, in addition to the classical fluorescence parameters, via the PAM fluorometer also the Chl fluorescence decrease ratio RFd (Fd/Fs), which, when measured at saturation irradiance is directly correlated to the net CO2 assimilation rate (P N) of leaves. This RFd-ratio can be determined from the Chl fluorescence induction kinetics measured with the PAM fluorometer using continuous saturating light (cSL) during 4–5 min. As the RFd-values are fast measurable indicators correlating with the photosynthetic activity of whole leaves, they should always be determined via the PAM fluorometer parallel to the other Chl fluorescence coefficients and ratios.

389 citations


Cites methods from "Chlorophyll Fluorescence Parameters..."

  • ...…several other Chl fluorescence parameters can be determined, e.g. Fv, Fv’, NF, and ∆F, which are used to calculate particular Chl fluorescence ratios applied as indicators of particular aspects of the photosynthetic apparatus (Lichtenthaler and Rinderle 1988b, Buschmann 1999, Roháček 2002)....

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Journal ArticleDOI
TL;DR: Questions about instruments, methods and applications based on chlorophyll a fluorescence, and the answers draw on knowledge from different Chl a Fluorescence analysis domains, yielding in several cases new insights.
Abstract: Using chlorophyll (Chl) a fluorescence many aspects of the photosynthetic apparatus can be studied, both in vitro and, noninvasively, in vivo. Complementary techniques can help to interpret changes in the Chl a fluorescence kinetics. Kalaji et al. (Photosynth Res 122:121–158, 2014a) addressed several questions about instruments, methods and applications based on Chl a fluorescence. Here, additional Chl a fluorescence-related topics are discussed again in a question and answer format. Examples are the effect of connectivity on photochemical quenching, the correction of F V /F M values for PSI fluorescence, the energy partitioning concept, the interpretation of the complementary area, probing the donor side of PSII, the assignment of bands of 77 K fluorescence emission spectra to fluorescence emitters, the relationship between prompt and delayed fluorescence, potential problems when sampling tree canopies, the use of fluorescence parameters in QTL studies, the use of Chl a fluorescence in biosensor applications and the application of neural network approaches for the analysis of fluorescence measurements. The answers draw on knowledge from different Chl a fluorescence analysis domains, yielding in several cases new insights.

384 citations


Cites background from "Chlorophyll Fluorescence Parameters..."

  • ...…several review papers have been published: Krause and Weis (1991), Dau (1994), Govindjee (1995), Rohácek and Barták (1999), Maxwell and Johnson (2000), Roháček (2002), Lazár (2006), Logan et al. (2007), Baker (2008), Henriques (2009), Allakhverdiev (2011), Gorbe and Calatayud (2012), Misra…...

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  • ...logical interpretation of Rfd values (Roháček 2002)....

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  • ...The fluorescence decrease from FP to FS depends on several factors that have not yet been characterized completely, complicating the physiological interpretation of Rfd values (Roháček 2002)....

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References
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Journal ArticleDOI
TL;DR: In this article, the quantum yield of non-cyclic electron transport was found to be directly proportional to the product of the photochemical fluorescence quenching (qQ) and the efficiency of excitation capture by open Photosystem II (PS II) reaction centres (Fv/Fm).

7,821 citations


"Chlorophyll Fluorescence Parameters..." refers background or methods in this paper

  • ...1): F F F F q q FS 0' V' V' P P V'( ) 1 , (19) from which the definition of the coefficient qP called the photochemical quenching of variable ChlF (Schreiber et al. 1986, Genty et al. 1989) follows: q F F F F F F F F F F F F F P S 0' V ' S 0' M ' 0 ' M ' S M ' 0 ' V ' 1 1 ....

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  • ...This well-known relationship (Genty et al. 1989) clarifies the dependence of 2 on four basic Chl FPs....

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  • ...They only differ by introduction of the substitution F F F   M S (Genty et al. 1989) and Eq....

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  • ...: the actual quantum yield of PS2 (non-cyclic) electron transport (Genty et al. 1989), efficiency of PS2 electron transport per quantum absorbed by PS2 complexes (ibidem), overall quantum yield of photochemical energy conversion in PS2 (Anonymous 1993), actual efficiency of energy conversion in PS2 (Björkman and Demmig-Adams 1995), quantum efficiency of PS2 photochemistry (Genty and Harbinson 1996), fraction of light absorbed in PS2 antennae utilised in PS2 photochemistry (Demmig-Adams et al....

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  • ...(37) This well-known relationship (Genty et al. 1989) clarifies the dependence of 2 on four basic Chl FPs....

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Journal ArticleDOI
TL;DR: An introduction for the novice into the methodology and applications of chlorophyll fluorescence is provided and a selection of examples are used to illustrate the types of information that fluorescence can provide.
Abstract: typically written from a biophysicist’s or a molecular plant physiologist’s point of view (Horton and Bowyer, Chlorophyll fluorescence analysis has become one of 1990; Krause and Weis, 1991; Govindjee, 1995). The aim the most powerful and widely used techniques avail- of this review is to provide a simple, practical guide to able to plant physiologists and ecophysiologists. This chlorophyll fluorescence for those beginners who are review aims to provide an introduction for the novice interested in applying the technique in both field and into the methodology and applications of chlorophyll laboratory situations. Whilst the principles behind the fluorescence. After a brief introduction into the theor- measurements will be discussed briefly, the emphasis will etical background of the technique, the methodology be on the applications and limitations of this technique and some of the technical pitfalls that can be encoun- in plant ecophysiology. tered are explained. A selection of examples is then used to illustrate the types of information that fluorescence can provide. The basis of chlorophyll fluorescence measurements

7,721 citations


"Chlorophyll Fluorescence Parameters..." refers background or methods in this paper

  • ...…the FV/FM ratio and/or maximum yield of primary photochemistry (Kitajima and Butler 1975), potential yield of PS2 photochemical reactions (Krause and Weis 1991), potential maximum PS2 quantum yield (Schreiber et al. 1995), and quantum efficiency of open PS2 centres (Maxwell and Johnson 2000)....

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  • ...1995), and quantum efficiency of open PS2 centres (Maxwell and Johnson 2000)....

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  • ...This technique has allowed an increased understanding of photochemical and non-photochemical processes occurring in thylakoid membranes of chloroplasts (for reviews, see Butler 1978, Walker 1987, Krause and Weis 1991, Govindjee 1995, Kramer and Crofts 1996, Lazár 1999, Maxwell and Johnson 2000)....

    [...]

  • ...The latter processes involve the pH-gradient build up, non-radiative energy dissipation (NRD) to heat within thylakoid membranes, LHC2 phosphorylation, photoinhibition of PS2 RCs, etc. (for reviews, see Krause and Weis 1991, Govindjee 1995, Maxwell and Johnson 2000)....

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Journal ArticleDOI
01 Jan 1991
TL;DR: Fluorescence as a Reaction Competing in the Deactivation of Excited Chlorophyll and the Origin of Fluorescence Emission.
Abstract: BIOPHYSICAL BASIS O F FLUORESCENCE EMISSION FROM CHLOROPLASTS . . . . .. . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . 314 Fluorescence as a Reaction Competing in the Deactivation of Excited Chlorophyll . . . . . . . . . . ... . . .. . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 314 Lifetimes of Fluorescence . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . 317 Origin of Fluorescence Emission . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32 1 Fluorescence of PS 11 and PS I at Ambient and Low Temperatures . . . . . . . . . . . . . . . . . . . 323 FLUORESCENCE INDUCTION AND PS II HETEROGENEITy 325 Fluorescence Transient from Fo to FM . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325 The FI Level and Inactive PS11 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .... .... . .. . . . .. . . . . . . . . . 326 Fluorescence Induction in High Ught .. . . . . . . . . 327 Rise in the Presence of DCMU and a/{3 Heterogeneity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 327 FLUORESCENCE QUENCHING 329 Resolution of Quenching Components . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . ........ . .. . . . . . . . . . . . . . . 330 Mechanism of Energy·Dependent Quenching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 3 1 Quenching Related t o State Transition . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . .. . ......... . . .. .. . . . . . . . 334 Photoinhibitory Quenching . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 334 Further Quenching Mechanisms . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 338 Physiological Aspects of Quenching . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . 338 CONCLUSIONS AND PERSPECTiVES 341

4,144 citations


"Chlorophyll Fluorescence Parameters..." refers background or methods in this paper

  • ...…by a pH-gradient build-up as well as activation of numerous regulatory mechanisms which provide for an effective utilisation of the excitation energy and cope with over-excitation (photoinhibition) or other kinds of injury caused to the thylakoid membrane (see Krause and Weis 1991, Horton 1996)....

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  • ...…called the FV/FM ratio and/or maximum yield of primary photochemistry (Kitajima and Butler 1975), potential yield of PS2 photochemical reactions (Krause and Weis 1991), potential maximum PS2 quantum yield (Schreiber et al. 1995), and quantum efficiency of open PS2 centres (Maxwell and Johnson…...

    [...]

  • ...The latter processes involve the pH-gradient build up, non-radiative energy dissipation (NRD) to heat within thylakoid membranes, LHC2 phosphorylation, photoinhibition of PS2 RCs, etc. (for reviews, see Krause and Weis 1991, Govindjee 1995, Maxwell and Johnson 2000)....

    [...]

  • ...The approximation presented by Oxborough and Baker (1997) presupposes that the PS2 photochemistry is actually trap-limited (cf. Krause and Weis 1991, Lavergne and Trissl 1995) and not limited by the rate of exciton transfer from a light-harvesting complex to PS2 RC (the diffusion limitation), as…...

    [...]

  • ...This technique has allowed an increased understanding of photochemical and non-photochemical processes occurring in thylakoid membranes of chloroplasts (for reviews, see Butler 1978, Walker 1987, Krause and Weis 1991, Govindjee 1995, Kramer and Crofts 1996, Lazár 1999, Maxwell and Johnson 2000)....

    [...]

Journal ArticleDOI
TL;DR: 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.
Abstract: 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.

2,671 citations


"Chlorophyll Fluorescence Parameters..." refers background or methods in this paper

  • ...1): F F F F q q FS 0' V' V' P P V'( ) 1 , (19) from which the definition of the coefficient qP called the photochemical quenching of variable ChlF (Schreiber et al. 1986, Genty et al. 1989) follows: q F F F F F F F F F F F F F P S 0' V ' S 0' M ' 0 ' M ' S M ' 0 ' V ' 1 1 ....

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  • ...(17) The definition for the coefficient of proportionality qN called the non-photochemical quenching of variable ChlF (Schreiber et al. 1986, van Kooten and Snel 1990) follows from Eq....

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  • ...The resolution of both principal types of the ChlF quenching was allowed by a saturation pulse method (Schreiber 1986, Schreiber et al. 1986), originally introduced as a “light-doubling” technique (Bradbury and Baker 1981, 1984)....

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  • ...19 results in another important relationship (cf. Schreiber et al. 1986): F F q q FS 0' P N V( )( ) 1 1 ....

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  • ...15, then (cf. Schreiber et al. 1986, Buschmann 1999): F F F F F q F FV' 0' V 0 V N 0 0' F q FV' N V( ) 1 ....

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Journal ArticleDOI
01 Apr 1987-Planta
TL;DR: Determinations of the photon yield of O2 evolution and the Fv/FM, 692 ratio can serve as excellent quantitative measures of photoinhibition of overall photosynthetic energy-conversion system and of photochemistry of photosystem II, respectively.
Abstract: Photon yields of oxygen evolution at saturating CO2 were determined for 44 species of vascular plants, representing widely diverse taxa, habitats, life forms and growth conditions. The photonyield values on the basis of absorbed light (φa) were remarkably constant among plants possessing the same pathway of photosynthetic CO2 fixation, provided the plants had not been subjected to environmental stress. The mean φa value ±SE for 37 C3 species was 0.106±0.001 O2·photon-1. The five C4 species exhibited lower photon yields and greater variation than the C3 species (φa=0.0692±0.004). The φa values for the two Crassulaceanacid-metabolism species were similar to those of C3 species. Leaf chlorophyll content had little influence on φa over the range found in normal, healthy leaves. Chlorophyll fluorescence characteristics at 77 K were determined for the same leaves as used for the photon-yield measurements. Considerable variation in fluorescence emission both at 692 nm and at 734 nm, was found 1) among the different species; 2) between the upper and lower surfaces of the same leaves; and 3) between sun and shade leaves of the same species. By contrast, the ratio of variable to maximum fluorescence emission at 692 nm (Fv/FM, 692) remained remarkably constant (The mean value for the C3 species was 0.832±0.004). High-light treatments of shade leaves resulted in a reduction in both φa and the Fv/FM, 692 ratio. The extent of the reductions increased with time of exposure to bright light. A linear relationship was obtained when φa was plotted against Fv/FM, 692. The results show that determinations of the photon yield of O2 evolution and the Fv/FM, 692 ratio can serve as excellent quantitative measures of photoinhibition of overall photosynthetic energy-conversion system and of photochemistry of photosystem II, respectively. This is especially valuable in field work where it is often impossible to obtain appropriate controls.

2,287 citations


"Chlorophyll Fluorescence Parameters..." refers background or methods in this paper

  • ...If the referred value P0 = 0.832 (Björkman and Demmig 1987) is used this ratio equals to 4.95....

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  • ...P0 is almost constant for many different plant species measured under the non-stressed conditions and equals to 0.832 0.004 (Björkman and Demmig 1987)....

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  • ...832 (Björkman and Demmig 1987) is used this ratio equals to 4....

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  • ...For healthy, non-stressed plants, it is mostly 0.8 P0 0.86 (Björkman and Demmig 1987) and therefore 0.14 F0/FM 0.2....

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It is demonstrated that all other Chl FPs can be determined using this quartet of parameters.