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

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 (_{P O}), the photochemical quenching of variable ChlF (q_P), the non-photochemical quenching of variable ChlF (q_N), and the relative change of minimum ChlF (q_O). _{P O} refers to the dark-adapted state of a thylakoid membrane, q_P, q_N and q_O 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 (q_CN), and the effective quantum yield of non-photochemical processes in PS2 (_N). New FPs, the total quenching of variable ChlF (q_TV) and the absolute quenching of ChlF (q_A) 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.).     

Ontogenetic approach to the assessment of plant resistance to prolonged stress using chlorophyll fluorescence induction method

The review deals with objective reasons that limit the use of chlorophyll fluorescence induction kinetics (Chl FIK) method in plant ecology. Based on the ontogenetic approach (analysis and comparison of the dynamics of the Chl fluorescence ratio F_p/F_s and physiological characteristics of plant leaves in ontogeny) possible criteria for the estimation of general plant resistance of photosynthetic apparatus (PSA) to prolonged stress affects are proposed. One of these criteria is the presence or absence of a steady-state phase in the dynamics of Chl fluorescence ratio F_p/F_s (or R_fd) of plant leaves after they stop growing. We also propose to use the duration of the steady-state phase and variability of Chl fluorescence ratios R_fd and F_p/F_s in this period for quantitative assessment of plant PSA resistance to prolonged stress during plant leaf ontogeny. The paper is dedicated to memory of Prof. F.Ya. Sid’ko.     

Chlorophyll fluorescence imaging of photosynthetic activity with the flash-lamp fluorescence imaging system

A flash-lamp chlorophyll (Chl) fluorescence imaging system (FL-FIS) is described that allows to screen and image the photosynthetic activity of several thousand leaf points (pixels) of intact leaves in a non-destructive way within a few seconds. This includes also the registration of several thousand leaf point images of the four natural fluorescence bands of plants in the blue (440 nm) and green (520 nm) regions as well as the red (near 690 nm) and far-red (near 740 nm) Chl fluorescence. The latest components of this Karlsruhe FL-FIS are presented as well as its advantage as compared to the classical single leaf point measurements where only the fluorescence information of one leaf point is sensed per each measurement. Moreover, using the conventional He-Ne-laser induced two-wavelengths Chl fluorometer LITWaF, we demonstrated that the photosynthetic activity of leaves can be determined measuring the Chl fluorescence decrease ratio, R_Fd (defined as Chl fluorescence decrease F_d from maximum to steady state fluorescence F_s:F_d/F_s), that is determined by the Chl fluorescence induction kinetics (Kautsky effect). The height of the values of the Chl fluorescence decrease ratio R_Fd is linearly correlated to the net photosynthetic CO₂ fixation rate P _N as is indicated here for sun and shade leaves of various trees that considerably differ in their P _N. Imaging the R_Fd-ratio of intact leaves permitted the detection of considerable gradients in photosynthetic capacity across the leaf area as well as the spatial heterogeneity and patchiness of photosynthetic quantum conversion within the control leaf and the stressed plants. The higher photosynthetic capacity of sun versus shade leaves was screened by Chl fluorescence imaging. Profile analysis of fluoresence signals (along a line across the leaf area) and histograms (the signal frequency distribution of the fluorescence information of all measured leaf pixels) of Chl fluorescence yield and Chl fluorescence ratios allow, with a high statistical significance, the quantification of the differences in photosynthetic activity between various areas of the leaf as well as between control leaves and water stressed leaves. The progressive uptake and transfer of the herbicide diuron via the petiole into the leaf of an intact plant and the concomitant loss of photosynthetic quantum conversion was followed with high precision by imaging the increase of the red Chl fluorescence F₆₉₀. Differences in the availability and absorption of soil nitrogen of crop plants can be documented via this flash-lamp fluorescence imaging technique by imaging the blue/red ratio image F₄₄₀/F₆₉₀, whereas differences in Chl content are detected by collecting images of the fluorescence ratio red/far-red, F₆₉₀/F₇₄₀.     

Photosynthesis performance in sweet almond [<Emphasis Type="Italic">Prunus dulcis</Emphasis> (Mill) D. Webb] exposed to supplemental UV-B radiation

Due to anthropogenic influences, solar UV-B irradiance at the earth’s surface is increasing. To determine the effects of enhanced UV-B radiation on photosynthetic characteristics of Prunus dulcis, two-year-old seedlings of the species were submitted to four levels of UV-B stress, namely 0 (UV-B_c), 4.42 (UV-B₁), 7.32 (UV-B₂) and 9.36 (UV-B₃) kJ m⁻² d⁻¹. Effects of UV-B stress on a range of chlorophyll (Chl) fluorescence parameters (FPs), Chl contents and photosynthetic gas-exchange parameters were investigated. UV-B stress promoted an increase in minimal fluorescence of dark-adapted state (F₀) and F₀/F_m, and a decrease in variable fluorescence (F_v, F_v/F_m, F_v/F₀ and F₀/F_m) due to its adverse effects on photosystem II (PSII) activity. No significant change was observed for maximal fluorescence of dark-adapted state (F_m). Enhanced UV-B radiation caused a significant inhibition of net photosynthetic rate (P _N) at UV-B₂ and UV-B₃ levels and this was accompanied by a reduction in stomatal conductance (g _s) and transpiration rate (E). The contents of Chl a, b, and total Chl content (a+b) were also significantly reduced at increased UV-B stress. In general, adverse UV-B effects became significant at the highest tested radiation dose 9.36 kJ m⁻² d⁻¹. The most sensitive indicators for UV-B stress were F_v/F₀, Chl a content and P _N. Significant P<0.05 alteration in these parameters was found indicating the drastic effect of UV-B radiation on P. dulcis.     

Chlorophyll fluorescence performance of sweet almond [Prunus dulcis (Miller) D. Webb] in response to salinity stress induced by NaCl

One-year old sweet almond (Prunus dulcis) seedlings were submitted to four levels of salt stress induced by NaCl, namely 0.3, 0.5, 0.7, and 1.0 S m⁻¹. Effects of salt stress on a range of chlorophyll (Chl) fluorescence parameters (Chl FPs) and Chl contents were investigated in order to establish an eco-physiological characterization of P. dulcis to salinity. Salt stress promoted an increase in F₀, F_s, and F₀/F_m and a decrease in F_m, F′_m, F_v/F_m, q_P, ΔF/F′_m, F_v/F₀, and UQF_(rel), in almost all Chl fluorescence yields (FY) and FPs due to its adverse effect on activity of photosystem 2. No significant changes were observed for quenchings q_N, NPQ, and q_N(rel). The contents of Chl a and b and their ratio were also significantly reduced at increased salt stress. In general, adverse salinity effects became significant when the electric conductivity of the nutrient solution (EC_n) exceeded 0.3 S m⁻¹. The most sensitive salt stress indicators were F_v/F₀ and Chl a content, and they are thus best used for early salt detection in P. dulcis. Monitoring of a simple Chl FY, such as F₀, also gave a good indication of induced salt stress due to the significant correlations observed between the different Chl FYs and FPs. Even essential Chl FYs, like F₀, F_m, F′_m, and F_s, and mutually independent Chl FPs, like F_v/F₀ and q_P, were strongly correlated with each other.     

Measurement of Differences in Red Chlorophyll Fluorescence and Photosynthetic Activity between Sun and Shade Leaves by Fluorescence Imaging

With a flash-lamp chlorophyll (Chl) fluorescence imaging system (FL-FIS) the photosynthetic activity of several thousand image points of intact shade and sun leaves of beech were screened in a non-destructive way within a few seconds. The photosynthetic activity was determined via imaging the Chl fluorescence at maximum F_p and steady state fluorescence F_s of the induction kinetics (Kautsky effect) and by a subsequent determination of the images of the fluorescence decrease ratio R_Fd and the ratio F_p/F_s. Both fluorescence ratios are linearly correlated to the photosynthetic CO₂ fixation rates. This imaging method permitted to detect the gradients in photosynthetic capacity and the patchiness of photosynthetic quantum conversion across the leaf. Sun leaves of beech showed a higher photosynthetic capacity and differential pigment ratios (Chl a/b and Chls/carotenoids) than shade leaves. Profile analysis and histogram of the Chl fluorescence yield and the Chl fluorescence ratios allow to quantify the differences in photosynthetic activity between different leaf parts and between sun and shade leaves with a high statistical significance.     

How to correctly determine the different chlorophyll fluorescence parameters and the chlorophyll fluorescence decrease ratio R<Subscript>Fd</Subscript> of leaves with the PAM fluorometer

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 F_v, F_m, F₀, F_m′, F_v′, NF, and ΔF, the Chl fluorescence ratios F_v/F_m, F_v/F₀, ΔF/F_m′, as well as the photochemical (q_P) and non-photochemical quenching coefficients (q_N, q_CN, and NPQ). q_N consists of three components (q_N = q_E + q_T + q_I), 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, q_N, q_CN, and NPQ increased in both leaf sides, but to a much higher extent at the lower compared to the upper leaf side. q_N was the best indicator for non-photochemical quenching even during a stronger HL-stress, whereas q_CN 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 R_Fd (F_d/F_s), which, when measured at saturation irradiance is directly correlated to the net CO₂ assimilation rate (_{P N}) of leaves. This R_Fd-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 R_Fd-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.     

Photochemical and Thermal Phases of Chlorophyll a Fluorescence

The measurement of variable chlorophyll (Chl) a fluorescence is widely used as a convenient and versatile tool in photosynthesis research. In many applications empirical correlations and simplified models of Chl a fluorescence are used with success. Nevertheless, variable Chl a fluorescence provides only indirect and complex image of processes occurring within photosynthetic membranes and such simplifications have only limited validity. In this review we elucidate some controversial and still unresolved questions about the origin and interpretation of the variable Chl a fluorescence induction and the proper use of variable Chl a fluorescence for studies of photochemical events in photosystem 2 (PS2). Although the major part of variable Chl a fluorescence reflects the photochemical closure of the PS2 reaction centers (RCs) and can be considered as a function of the redox state of the primary acceptor Q_A, up to 50 % of the change in the Chl a fluorescence yield can be of secondary, nonphotochemical origin. We review the possible sources of the inherent heterogeneity in the origin of variable Chl a fluorescence. We also comment on the practical implications this bears for the use of variable Chl a fluorescence. This revised version was published online in August 2006 with corrections to the Cover Date.     

外源甜菜碱对盐胁迫下枸杞光合功能的改善

研究了外源甜菜碱对盐胁迫下枸杞扦插苗叶片光合功能的影响。结果表明,外源甜菜碱能使盐胁迫下的枸杞叶片叶绿素荧光动力学参数Fo、Fm、Fv、Fv／Fm、Fm／Fo和Fv／Fo增高,使光合色素叶绿素a(Chla)、叶绿素b(Chlb)和类胡萝卜素(Car)含量明显增加,叶绿素a与b的比值(Chla／Chlb)升高,类胡萝卜素与叶绿素的比值(Car／Chl)降低,说明外源甜菜碱有利于植物对光能的捕获、吸收、传递和转换,提高叶片的光合活性,降低盐胁迫对植物的抑制作用。     

Photosynthetic activity of poikilochlorophyllous desiccation tolerant plant <Emphasis Type="Italic">Reaumuria soongorica</Emphasis> during dehydration and re-hydration

Diurnal patterns of gas exchange and chlorophyll (Chl) fluorescence parameters of photosystem 2 (PS2) as well as Chl content were analyzed in Reaumuria soongorica (Pall.) Maxim., a perennial semi-shrub during dehydration and rehydration. The net photosynthetic rate (P _N), maximum photochemical efficiency of PS2 (variable to maximum fluorescence ratio, F_v/F_m), quantum efficiency of non-cyclic electron transport of PS2, and Chl content decreased, but non-photochemical quenching of fluorescence and carotenoid content increased in stems with the increasing of drought stress. 6 d after re-hydration, new leaves budded from stems. In the re-watered plants, the chloroplast function was restored and Chl a fluorescence returned to a similar level as in the control plants. This improved hydraulic adjustment in plant triggered a positive effect on ion flow in the tissues and increased shoot electrical admittance. Thus R. soongorica plants are able to sustain drought stress through leaf abscission and keep part of Chl content in stems.     

Satiation-dependent, intra-cohort variations in prey size selection of young roach (Rutilus rutilus)

The tolerance to freezing and thawing of Leucodon sciuroides, a moss growing in mountainous areas of the Mediterranean (south-east Spain), was investigated by means of CO₂ gas exchange, modulated chlorophyll (Chl) a fluorescence and pigment analysis by high-performance liquid chromatography. Evidence is presented for freezing-induced decreases in CO₂ fixation that enhance non-radiative dissipation of absorbed light energy, a process which protects the photosynthetic apparatus. The photosynthetic apparatus of L. sciuroides remained fully recuperable after freezing, as indicated by the recovery of photosynthetic CO₂ fixation and Chl fluorescence parameters to pre-freezing values during thawing. The rapid recovery of photosynthesis activity during thawing in L. sciuroides suggests that this moss is capable of tolerating freeze-thaw cycles in a manner similar to that found at higher latitudes or in the Antarctic. The resistance of the photosynthetic apparatus of this moss to freezing might be achieved, at least partially, through the employment of dissipative pathways, such as non-radiative dissipation of absorbed light energy. Received: 4 June 1998 / Accepted: 15 February 1999     

A novel insight into the regulation of light-independent chlorophyll biosynthesis in <Emphasis Type="Italic">Larix decidua</Emphasis> and <Emphasis Type="Italic">Picea abies</Emphasis> seedlings

Light-independent chlorophyll (Chl) biosynthesis is a prerequisite for the assembly of photosynthetic pigment–protein complexes in the dark. Dark-grown Larix decidua Mill. seedlings synthesize Chl only in the early developmental stages and their Chl level rapidly declines during the subsequent development. Our analysis of the key regulatory steps in Chl biosynthesis revealed that etiolation of initially green dark-grown larch cotyledons is connected with decreasing content of glutamyl-tRNA reductase and reduced 5-aminolevulinic acid synthesizing capacity. The level of the Chl precursor protochlorophyllide also declined in the developing larch cotyledons. Although the genes chlL, chlN and chlB encoding subunits of the light-independent protochlorophyllide oxidoreductase were constitutively expressed in the larch seedlings, the accumulation of the ChlB subunit was developmentally regulated and ChlB content decreased in the fully developed cotyledons. The efficiency of chlB RNA-editing was also reduced in the mature dark-grown larch seedlings. In contrast to larch, dark-grown seedlings of Picea abies (L.) Karst. accumulate Chl throughout their whole development and show a different control of ChlB expression. Analysis of the plastid ultrastructure, photosynthetic proteins by Western blotting and photosynthetic parameters by gas exchange and Chl fluorescence measurements provide additional experimental proofs for differences between dark and light Chl biosynthesis in spruce and larch seedlings.     

Effects of Season,Needle Age,and Elevated Atmospheric CO2 on Chlorophyll Fluorescence Parameters and Needle Nitrogen Concentration in Scots Pine (Pinus sylvestris)

Six-year-old Scots pine (Pinus sylvestris L.) seedlings were grown in open top chambers (OTCs) at ambient (AC) or elevated (ambient + 400 µmol mol^–1; EC) CO₂ concentration for three years (1996–1998). Chlorophyll (Chl) a fluorescence of current and one-year-old needles was measured in the field at two-weekly intervals in the period July–October 1998. In addition, Chl, carbon (C), and nitrogen (N) concentrations in both needle age classes were determined monthly during the same period. Chl fluorescence parameters were not significantly affected by EC, suggesting there was no response of the light reactions and the photochemical efficiency of photosystem 2. Chl concentrations were not significantly different but a reduced N concentration was observed in needles of EC treatment. Significant differences between needle age classes were observed for all parameters, but were most apparent under EC and toward the end of the growing season, possibly due to an acclimation process. As a result, significant interactions between CO₂ treatment, needle age class, and season were found. This study emphasizes the importance of repeated measures including different leaf/needle age classes to assess the photosynthetic response of trees under EC.     

Low Temperature Tolerance of Tobacco Plants Transformed to Accumulate Proline,Fructans, or Glycine Betaine. Variable Chlorophyll Fluorescence Evidence

Tobacco (Nicotiana tabacum L.) has been transformed to accumulate different compatible solutes (proline, fructans, or glycine betaine) in order to improve its tolerance to abiotic stress. Photosynthetic activity of wild Type (wt) and transformed tobacco plants before and after freezing stress was studied by measuring chlorophyll (Chl) fluorescence. The JIP test of Chl fluorescence induction was used to analyze in details the functional activity of photosystem 2. No significant differences were found among wild Type and transgenic plants after 12 h of freezing. Both plant Types maintained the same values of the measured parameters [F_V/F_M, PI(CS_M), ABS/RC, TR₀/RC, ET/RC] after recovery of stress. The studied Chl fluorescence parameters decreased only for the wild Type plants, stressed for 24 h at –2 °C. The strong inhibition of photosynthetic reactions in the wt plant after 24 h of freezing could not be restored. The evaluated parameters of transgenic plants did not change significantly after 24 h at –2 °C and successfully survived freezing stress.     

Effects of foliar application of nitrogen on the photosynthetic performance and growth of two fescue cultivars under heat stress

The effects of nitrogen fertilization on the growth, photosynthetic pigment contents, gas exchange, and chlorophyll (Chl) fluorescence parameters in two tall fescue cultivars (Festuca arundinacea cv. Barlexas and Crossfire II) were investigated under heat stress at 38/30 °C (day/night) for two weeks. Shoot growth rate of two tall fescue cultivars declined significantly under heat stress, and N supply can improved the growth rates, especially for the Barlexas. Chl content, leaf net photosynthetic rate, stomatal conductance, water use efficiency, and the maximal efficiency of photosystem 2 photochemistry (F_v/F_m) also decreased less under heat stress by N supply, especially in Crossfire II. Moreover, cultivar variations in photosynthetic performance were associated with their different response to heat stress and nitrogen fertilization, which were evidenced by shoot growth rate and photosynthetic pigment contents.     

Differential protection of photosynthetic capacity in trehalose-and lea protein-producing transgenic plants under abiotic stresses

We previously demonstrated that both trehalose and LEA protein protect plants from damage by drought, salt, and heat. Here, we compared their effectiveness in preserving photosynthetic capacity under those abiotic stresses. Upon dehydration, the Pmax (maximal photosynthetic rate) of O₂ evolution decreased similarly in both nontransformants andotsA plants. Contrastingly, Pmax was maintained at a considerably higher level inCaLEA6 plants. However, no significant differences in Chl fluorescence parameters were observed between transformants and nontransformants. Under salinity stress,CaLEA6 plants were also better thanotsA plants in terms of their values for Pmax, photochemical efficiency of PSII(Fv/Fm), and photochemical quenching (qP). After heat bothotsA andCaLEA6 plants maintained a higher Pmax as well as more favorable Chl fluorescence parameters, although the latter transformant performed slightly better overall. Therefore, despite the comparable effectiveness of trehalose and LEA protein in enhancing tolerance against those abiotic stresses, they confer differential protection in maintaining photosynthetic capacity. Compared with trehalose, the CaLEA6 protein appears to be a more universal and effective agent under those stresses.     

Photosynthesis in different types of transgenic tobacco plants with elevated cytokinin content

Photosynthetic parameters were compared in three types of transgenic tobacco plants: ipt-transgenic plants with slightly elevated endogenous cytokinin (CK) content, Pssu-ipt-transgenic plants with markedly increased CK content, and zmp-transgenic plants with slightly elevated CK content accompanied by elevated auxin content. Slightly increased CK content promoted net photosynthetic rate (P_N) in both ipt- and zmp-transgenic plants, and chlorophyll (Chl) and carotenoid contents in zmp-transgenic plants. Morphology, growth characteristics, stomatal conductance, and parameters of Chl a fluorescence kinetics were similar in control and transgenic plants with slightly higher CK content. No significant effect of increased level of endogenous auxin (indole-3-acetic acid) on development of zmp-transgenic plants and measured parameters was found. Pssu-ipt-transgenic plants with highly increased CK content revealed suppressed root development, wilting of plants, and depression of P_N and stomatal conductance; however, Chl content was slightly increased and parameters of Chl a fluorescence kinetics did not indicate damage to photosynthetic apparatus. This revised version was published online in July 2006 with corrections to the Cover Date.     

Frequently asked questions about chlorophyll fluorescence,the sequel

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.     

Mechanisms of non-photochemical chlorophyll fluorescence quenching in higher plants

The excitation energy of pigment molecules in photosynthetic antennae systems is utilised by photochemistry, partly it is thermally dissipated, and partly it is emitted as fluorescence. Changes in the quantum yield of chlorophyll (Chl) fluorescence reflect the changes in quantum yield of photochemical reaction and thermal dissipation of the excitation energy. Decrease of the Chl fluorescence quantum yield is called the Chl fluorescence quenching. The decrease of the quantum yield that is accompanied by photochemical reactions has been termed the photochemical quenching, and the decrease accompanied by thermal dissipation of the excitation energy is called the non-photochemical quenching. This review deals with mechanisms of the non-photochemical quenching.