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Article

Light Energy Use Efficiency in Photosystem II of Tomato Is Related to Leaf Age and Light Intensity

by
Julietta Moustaka
1,†,
Ilektra Sperdouli
2 and
Michael Moustakas
1,*
1
Department of Botany, Aristotle University of Thessaloniki, 54124 Thessaloniki, Greece
2
Institute of Plant Breeding and Genetic Resources, Hellenic Agricultural Organisation-Demeter (ELGO-Demeter), 57001 Thessaloniki, Greece
*
Author to whom correspondence should be addressed.
†
Current address: Department of Food Science, Aarhus University, 8200 Aarhus, Denmark.
Crops 2024, 4(4), 623-635; https://doi.org/10.3390/crops4040043
Submission received: 10 September 2024 / Revised: 10 October 2024 / Accepted: 7 November 2024 / Published: 11 November 2024
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Abstract

:
The fundamental key to increase photosynthetic efficiency of crop plants lies in optimizing the light energy use efficiency. In our study, we used tomato to evaluate the allocation of absorbed light energy in young and mature leaves, and to estimate if the extent of photoinhibition and photoprotection can be affected by the leaf age. A reduced efficiency of the oxygen-evolving complex, in young leaves compared to mature ones, resulted in a donor-side photoinhibition, as judged from the significantly lower Fv/Fm ratio, in young leaves. The detected increased 1O2 production in young leaves was probably due to a donor-side photoinhibition. The effective quantum yield of photosystem II (PSII) photochemistry (ΦPSII), at low light intensity (LLI, 426 μmol photons m−2 s−1), was significantly lower in young compared to mature leaves. Moreover, the non-significant increase in non-photochemical energy loss in PSII (ΦNPQ) could not counteract the decreased ΦPSII, and as a result the non-regulated energy loss in PSII (ΦNO) increased in young leaves, compared to mature ones. The significantly lower ΦPSII in young leaves can be attributed to the increased reactive oxygen species (ROS) creation that diminished the efficiency of the open PSII reaction centers (Fv’/Fm’), but without having any impact on the fraction of the open reaction centers. The reduced excess excitation energy, in mature leaves compared to young ones, at LLI, also revealed an enhanced PSII efficiency of mature leaves. However, there was almost no difference in the light energy use efficiency between young and mature leaves at the high light intensity (HLI, 1000 μmol photons m−2 s−1). The ability of mature tomato leaves to constrain photoinhibition is possible related to an enhanced photosynthetic function and a better growth rate. We concluded that the light energy use efficiency in tomato leaves is influenced by both the leaf age and the light intensity. Furthermore, the degrees of photoinhibition and photoprotection are related to the leaf developmental stage.

1. Introduction

By the process of photosynthesis, solar energy is converted into chemical energy, which is the energy for the synthesis of all the essential organic molecules that maintain the life of all living organisms on Earth [1,2,3,4]. Leaves are the principal photosynthetic organs of plants and above 90% of crop dry matter originates from leaf photosynthesis [4,5]. The light-harvesting complexes (LHCII) of photosystem II (PSII), also called antennae, transfer the absorbed light energy to the reaction centers (RCs) of PSII. PSII is a large pigment–protein complex that consists of a water-splitting system (oxygen-evolving complex, OEC), an LHCII, and the RCs [6]. At the OEC of PSII, the oxidation of water results in O2, H+, and e− [6].
Light absorption results in singlet-excited state chlorophyll molecules (1Chl*), which can be de-excited either (i) by safely dissipating energy as heat (referred as non-photochemical quenching, NPQ); (ii) by transferring the energy to another molecule and finally at the RC that can be de-excited by losing an electron, which is called photochemistry (referred to as qp); or (iii) by re-emitting light through fluorescence [6,7,8]. By translating the information of the re-emitted chlorophyll a fluorescence, accurate evidence about the status of the photosynthetic apparatus, and especially of PSII function, can be obtained [9,10,11,12,13]. One of the most efficient non-invasive techniques for the evaluation of photosynthetic performance, and abiotic or biotic stress tolerance monitoring, is chlorophyll-a fluorescence imaging, which can also estimate spatiotemporal variations in photosynthetic function across the whole-leaf area [10,12,13,14,15,16].
Heterogeneity of photosynthetic function between leaves of the same plant has been generally reported for different plant species [17]. Photosynthetic ability is influenced by the leaf structural differences, e.g., the mesophyll thickness and the chloroplast’s space [17,18]. When young leaves are exposed to light, only a portion of the captured light energy can be utilized in photochemical reactions, which means that excessive excited energy is produced [19,20]. Excess excitation energy in PSII can be dissipated as harmless heat through the protective mechanism of NPQ [7], or if it is not, it can generate reactive oxygen species (ROS) [8,21,22,23]. Thus, when the absorbed light energy is in excess of what can be handled by the electron transport chain capacity, an increased probability of ROS generation occurs [6,21,22,23].
During the light reactions of photosynthesis, ROS are constantly formed, but they are removed by the antioxidant mechanisms [23,24,25,26]. The ROS that are formed are the superoxide anion radical (O2•−), hydrogen peroxide (H2O2), and singlet-excited oxygen (1O2) [23,24,25,26]. Light energy capture by the photosynthetic pigments and photosynthetic performance are facilitated by leaf growth [27]. In young leaves, the light energy capture ability develops earlier than the CO2 assimilation ability [20,28], and thus ROS generation is enhanced when CO2 assimilation is inhibited [24,29].
Photosynthesis in mature leaves of Nerium oleander and Cercis siliquastrum was superior compared to young leaves, with a higher electron transport rate and better CO2 assimilation [30,31]. In contrast, there was no difference in photosynthesis between young and mature leaves in Phlomis fruticosa [30]. Diverse leaf developmental stages may present differential responses to drought stress, due to a differential accumulation of metabolites, and may show differential responses of PSII photochemistry to a water deficit, not following a dose-dependent reaction, but a hormetic response [32,33]. In Arabidopsis thaliana exposed to drought stress, PSII function in young leaves was found to acclimate better than mature ones, to a water deficit [34]. Ozone stress response of photosynthesis in Betula pubescens was found to depend on leaf age [35]. Young leaves manage to be more resilient to a water deficit than mature leaves [20,34,36], by accumulating higher concentrations of proline, sugars, and flavonoids, and showing lower oxidative damage and less excitation pressure, which motivates the acclimation of PSII photochemistry to a water deficit [32,37]. Young leaves also maintained their capability to recommence leaf expansion after water was resupplied [27,38].
Light is fundamental for photosynthesis, and low light often limits photosynthesis, but an excessive amount of light can possibly pose an injury [39]. Young leaves and old senescing leaves were found to be the most sensitive to light [40]. Low light intensities frequently restrict plant growth, also downgrading plants’ nutritional value, and being a limiting factor for crop production [41]. Crop monitoring for photosynthetic performance by chlorophyll-a fluorescence imaging techniques permits not only crop performance evaluation but also the early pre-symptomatic examination of the crop physiological status, allowing for the prevention of damage and major yield losses [3,13].
Tomato (Solanum lycopersicum L.) is the second-most important vegetable after potato, due to its high economical value and its significance in nutrition and health [42]. It is recognized as a rich source of compounds for the prevention of cardiovascular diseases and certain kinds of cancer [43]. It contains many healthy beneficial compounds such as flavonoids, vitamin E, ascorbic acid (vitamin C), β-carotene, and lycopene [43,44]. Due to its significance as a food source, extensive research has been conducted to enhance tomato crop productivity, improve its fruit quality, and increase its biotic and abiotic stress resistance [45].
Taking into account the bibliographic data that describe differential photosynthetic performance and photoprotective mechanisms between different leaf developmental stages of the same plant, we hypothesized that the light energy use efficiency in tomato leaves will be influenced by the leaf age, and the light intensity. We also evaluated if there is an interaction between the leaf age and the light intensity that could influence tomato PSII function. In addition, we examined the perspective of obtaining more information about how the extent of photoinhibition and photoprotection can be affected by the leaf developmental stage.

2. Materials and Methods

2.1. Plant Material

Tomato (Lycopersicon esculentum Mill. cv ecstasy) plants, with leaf developmental stage 15 according to the BBCH numerical scale, a system for uniform coding of growth stages [46], were obtained from a marketplace. Afterwards, plants were transported to a growth chamber with a 25 ± 1/20 ± 1 °C day/night temperature, a 14 h day/night photoperiod offered by a photosynthetic photon flux density (PPFD) of 420 ± 10 μmol quanta m−2 s−1, and a relative humidity of 65 ± 5/75 ± 5%, day/night.

2.2. Chlorophyll Fluorescence Imaging Analysis

The Imaging-PAM Fluorometer M-Series (Heinz Walz GmbH, Effeltrich, Germany) was used to estimate PSII function as described in detail previously [47]. Young (still developing) and mature (fully mature) tomato leaves (Figure S1) were dark-adapted for 30 min, and then chlorophyll fluorescence measurements were performed. In each leaf, 7 areas of interest (AOIs) were chosen to be calculated, so as to cover the whole-leaf area. Two actinic light (AL) intensities were used: 426 μmol photons m−2 s−1 (low light intensity, LLI) and 1000 μmol photons m−2 s−1 (high light intensity, HLI). The measured chlorophyll fluorescence parameters that are described in detail in Supplementary Table S1 were computed using the Win software V2.41a (Heinz Walz GmbH, Effeltrich, Germany). Leaf color-coded images of the photosynthetic parameters were also obtained at both AL intensities.

2.3. Statistical Analysis

The statistical analysis was performed with R software (version 4.3.1 R Core Team, 2023). A two-way ANOVA was performed for each parameter with leaf age (young or mature) and light intensity (426 μmol photons m−2 s−1 or 1000 μmol photons m−2 s−1) as factors, followed by a post hoc analysis with Tukey’s honest significant difference method with the R package ‘multcomp’. Values were considered significantly different at p < 0.05.

3. Results

3.1. The Impact of Leaf Age on the Efficiency of the Oxygen-Evolving Complex and the Maximum Efficiency of Photosystem II Photochemistry

The efficiency of the oxygen-evolving complex (OEC) in young leaves was significantly lower (−15%) compared to mature leaves (Figure 1a). Similarly, the maximum efficiency of PSII photochemistry (Fv/Fm), in dark-adapted tomato leaves, was also significantly lower (−3%) in young leaves compared to mature ones (Figure 1b).

3.2. Light Energy Use Efficiency in Young and Mature Leaves

We estimated the light energy partitioning at PSII, which is either distributed to photochemistry (ΦPSII), is dissipated as heat (ΦNPQ), or is lost by a non-regulatory way (ΦNO) [47]. The sum of all is equal to 1 [48]. Actinic light was applied to young and mature tomato leaves at two different intensities: 426 μmol photons m−2 s−1 (LLI) and 1000 μmol photons m−2 s−1 (HLI).
The quantum yield for photochemistry (ΦPSII) was affected by both leaf age and light intensity (Supplementary Table S2). ΦPSII was significantly lower (−7%) in young leaves, compared to mature leaves, at the LLI, while at the HLI, there was no significant difference (Figure 2a). In contrast to ΦPSII, the regulated non-photochemical energy loss in PSII (ΦNPQ) was only affected by light intensity (Supplementary Table S2) and was non-significantly higher in young leaves, compared to mature leaves, at the LLI, and also at the HLI, there was no significant difference (Figure 2b).
The yield of non-regulated energy loss in PSII (ΦNO) was affected by both factors (light intensity and leaf age), and it was significantly higher (+8%) in young leaves, compared to mature leaves, at both LLI and HLI (Figure 2c).

3.3. Impact of Leaf Age on Fraction of Open PSII Reaction Centers and Their Efficiency

The redox state of quinone A (QA), which also indicates the fraction of open PSII reaction centers (RCs) (qp), did not differ in young leaves, compared to mature leaves, at both LLI and HLI (Figure 3a).
Both light intensity and leaf age significantly affected the efficiency of the open PSII RCs (Fv’/Fm’). Mature tomato leaves exhibited significantly higher (Fv’/Fm’) (+5%), compared to young leaves, at both LLI and HLI (Figure 3b).

3.4. Electron Transport Efficiency and Photoprotective Heat Dissipation in Young and Mature Leaves

The electron transport rate (ETR) of PSII, in both young and mature leaves, increased by light intensity. At low light intensity (LLI), young leaves exhibited a significantly lower (−7%) ETR, compared to mature leaves, while at high light intensity (HLI), there was no significant difference (Figure 4a). The non-photochemical quenching (NPQ) did not differ in young leaves, compared to mature leaves, at both LLI and HLI (Figure 4b).

3.5. The Impact of Leaf Age on the Excess Excitation Energy at PSII and on the Excitation Pressure at PSII

Young tomato leaves at LLI exhibited significantly higher (+7%) excess excitation energy (EXC), compared to mature leaves, while at HLI, the EXC increased in both leaf types, but did not differ between them (Figure 5a).
The excitation pressure at PSII (1 − qL) did not differ in young leaves, compared to mature leaves, at both LLI and HLI (Figure 5b).

3.6. The Spatiotemporal Heterogeneity of PSII Photochemistry in Young and Mature Leaves

Figure 6 shows representative pseudocolor-coded pictures of the whole area of tomato leaves, for the parameters Fv/Fm (in dark-adapted leaves), and of ΦPSII, ΦNPQ, and qp, at 426 μmol photons m−2 s−1 (low light intensity, LLI) and at 1000 μmol photons m−2 s−1 (high light intensity, HLI), from young and mature leaves. A higher spatial heterogeneity, for the parameters ΦPSII and ΦNPQ, was noticed at LLI, while at HLI, a higher spatial heterogeneity was observed for the parameter qp (Figure 6). It looks like the increased dissipation of the absorbed light energy as heat (ΦNPQ) at the HLI, which drastically decreased ΦPSII, reduced the spatial PSII heterogeneity of ΦPSII and ΦNPQ, compared to LLI (Figure 6). In contrast, the heterogenous decreased fraction of open PSII reaction centers at HLI increased the spatial heterogeneity of qp (Figure 6).

4. Discussion

In the light reactions of photosynthesis, all plants use the light energy to initiate primary production, and thus it is generally recognized that the light energy use efficiency controls crop yields [49,50,51,52,53,54,55]. The absorbed light energy that is driven to photochemistry, also known as effective quantum yield of PSII photochemistry (ΦPSII), was significantly lower in young leaves, compared to mature leaves, at the LLI (Figure 2a). The accompanied non-significant increase in non-photochemical energy loss in PSII (ΦNPQ) in the young leaves (Figure 2b) could not counteract the decreased ΦPSII, and as a result the non-regulated energy loss in PSII (ΦNO) increased in young leaves, compared to mature leaves (Figure 2c). The increased ΦNO suggests an increased ROS creation, since ΦNO is related to the amount of 1O2 production [56,57,58]. 1O2 is produced when 1Chl* is not quenched, and through an intersystem crossing, the triplet chlorophyll state (3Chl*) is developed, leading to the generation of 1O2 [7,23,59,60]. The form of molecular oxygen in an excited state, i.e., 1O2, attacks and oxidizes electron-rich molecules, such as nucleic acids, lipids, and proteins [61,62,63], and consequently it is a harmful ROS molecule [64]. ROS production can be avoided by attenuating 1Chl* through the mechanism of NPQ, or by quenching 3Chl* [8,24,25,59]. Still, despite these protective mechanisms, detrimental 1O2 are created, related to the physiological phenomenon of photoinhibition [64,65,66,67,68,69].
Young tomato leaves did not show any enhancement of the photoprotective mechanism of NPQ (Figure 4b), to counteract the increased ROS creation that they presented, compared to mature ones [21,24,60,70]. As a consequence, 1O2 (Figure 2c) was generated, suggesting an increased ROS production in young leaves compared to mature ones. This is correlated with the decreased CO2 assimilation ability of young leaves compared to mature ones [20,28], which has been shown to result in enhanced ROS generation [24,29]. Our data of the NPQ levels in young and mature leaves, which were the same under both LLI and HLI (Figure 4b), are in agreement with the NPQ levels observed in young and mature wild-type Arabidopsis thaliana leaves [40].
An amplified ROS creation constrains the repair of photodamaged PSII RCs and mainly the de novo synthesis of the D1 protein [71,72], thus diminishing the efficiency of PSII RCs (Fv’/Fm’) as was observed in young leaves (Figure 3b). A reduced ΦPSII can be attributed (i) to a lower fraction of open PSII RCs (qp) or (ii) to a diminished efficiency of RCs (Fv’/Fm’) [73]. The reduced ΦPSII in young leaves compared to mature ones was ascribed to the reduced efficiency of the RCs (Fv’/Fm’) (Figure 3b) since the percentage of open PSII RCs (qp) remained the same with mature leaves (Figure 3a). Thus, the diminished efficiency of PSII RCs (Fv’/Fm’) in young leaves caused a reduced ETR (Figure 4a) and a reduced ΦPSII (Figure 2a), compared to mature ones.
Photoinhibition, also called photoinactivation, occurs when the excess energy absorbed by the leaves is neither consumed for photosynthesis nor dissipated safely by NPQ, or when the primary cause is the excitation of Mn in the OEC by photons [74,75,76,77,78,79,80]. In young leaves, a reduced efficiency of the OEC, compared to mature leaves, was observed, as judged from the ratio Fv/Fo [81,82,83,84,85,86,87]. The superior efficiency of the OEC in mature leaves, compared to young ones (Figure 1a), was in accordance with the corresponding enhanced maximum efficiency of PSII photochemistry (Fv/Fm) in mature leaves (Figure 1b). A lower OEC efficiency leads to a decrease in Fv/Fm [88,89], as was also observed in our experiment. The donor-side photoinhibition is regularly connected with ROS production [58,90]. The detected amplified 1O2 production in young leaves, compared to mature leaves (Figure 2c), was probably attributed to a donor-side photoinhibition (Figure 1b), due to the decreased efficiency of the OEC (Figure 1a). Photoinhibition has been shown to be correlated with a reduced efficiency of the OEC [91,92,93,94]. If the OEC does not appropriately reduce the primary electron donor, then it can cause destructive oxidations in PSII [94].
Plants activate numerous mechanisms that permit efficient acclimation to HLI [95,96]. Although HLI can boost photosynthesis, it can also harm photosynthetic procedures, especially together with other environmental stresses [97]. Plant acclimation to environmental stresses usually coincides with a high photosynthetic ability in HLI and more effective light harvesting and effective quantum yield (ΦPSII) under LLI [97]. Mature tomato leaves without showing a significant superior photosynthetic ability at HLI, compared to young ones, displayed a significant more effective quantum yield (ΦPSII) under LLI (Figure 2a and Figure 6).
Light, despite being an essential resource for photosynthesis, can trigger a decrease in photosynthetic function linked mainly with PSII photoinactivation [78]. Fv/Fm is typically correlated with photoinhibition and associated with a better growth rate [97]. The ability of mature tomato leaves to constrain photoinhibition (Figure 1b) is possibly related to an enhanced photosynthetic function and a better leaf growth rate. An increased ability of photosynthetic function reduces susceptibility to photodamage [96]. The reduced excess excitation energy (EXC), in mature leaves compared to young ones at LLI (Figure 5a), reveals an enhanced PSII efficiency of mature leaves. Leaves of different developmental stages at different sites in the canopy have anatomical, structural, and metabolic differences, which affect photosynthetic activity [98,99,100]. In environmental stress studies, it is crucial to consider the leaf developmental stage that affects photosynthetic function and compare leaves of the same developmental stage [17,27,34,101,102,103]. Our data (Table S2) show that leaf age and light intensity play crucial roles in determining PSII responses of tomato. However, the lack of significance in the interaction leaf age × light intensity is indicating an additive rather than an interactive relationship.
The evaluation of PSII functionality at the level of the whole leaf revealed a higher spatial heterogeneity at LLI for the parameters ΦPSII and ΦNPQ, while at HLI, this was the case for the parameter qp (Figure 6). Probably, the increased dissipation of the absorbed light energy as heat (ΦNPQ) at HLI, which also decreased ΦPSII significantly, in both leaf types, reduced the spatial heterogeneity. The observed spatial heterogeneity of PSII function in both tomato leaf types suggests that stomatal function and water potential differ in different regions of the leaf, contributing to spatial differences in PSII functionality and reflecting different zones of mesophyll development and leaf anatomy [104,105]. Improving photosynthesis is a key task for plant scientists, especially specified by the worldwide rising need for nutrition [106,107,108]. The fundamental key to increase photosynthetic efficiency lies in optimizing the allocation of absorbed light energy [34,109].

5. Conclusions

We conclude that the light energy use efficiency in tomato leaves is influenced by both the leaf age and the light intensity. Furthermore, the degree of photoinhibition and photoprotection are related to the leaf developmental stage. Mature leaves used the absorbed light energy more efficiently for ETR than young leaves, possessing superior efficiency of the oxygen-evolving complex, higher effective quantum yield, and reduced excess excitation energy at the low light intensity, but almost without any difference to young leaves at the high light intensity. We would like to highlight that our results support the understanding that tomato leaf age impacts photosynthetic efficiency and that light intensity consistently modulates this effect without introducing further complexity through interaction. Based on our results, we would like to recommend that in agricultural studies, it is essential to consider the leaf developmental stage and always compare leaves of the same developmental stage, examined under the same quantity and quality of light. Distinct leaf developmental stages may present differential responses to biotic and abiotic stresses, due to a differential accumulation of metabolites, and may show dissimilar physiological responses.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/crops4040043/s1, Figure S1. Tomato (Lycopersicon esculentum Mill. cv ecstasy) plant with arrows showing young and mature leaves that were measured. Table S1: Definitions of the chlorophyll fluorescence parameters used in the experiments. Table S2. Analysis of variance for parameters ΦPSII, ΦNPQ, ΦNO, and Fv’/Fm’ with factors leaf age, light intensity, and their interaction.

Author Contributions

Conceptualization, M.M.; methodology, J.M., I.S. and M.M.; validation, J.M. and M.M.; formal analysis, J.M., I.S. and M.M.; investigation, I.S. and M.M.; resources, M.M.; data curation, J.M., I.S. and M.M.; writing—original draft preparation, J.M. and M.M.; writing—review and editing, J.M., I.S. and M.M.; visualization, J.M., I.S. and M.M.; supervision, M.M.; project administration, M.M. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Data Availability Statement

The data presented in this study are available in this article.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. The efficiency of the oxygen-evolving complex (Fv/Fo) (a), and the maximum efficiency of PSII photochemistry (Fv/Fm) (b), in young and mature tomato leaves after 30 min of dark adaptation. Standard deviations (SDs) are shown by bars. Significant differences are shown by different lower-case letters (p < 0.05). Eight to ten plants were measured from each treatment (n = 8–10).
Figure 1. The efficiency of the oxygen-evolving complex (Fv/Fo) (a), and the maximum efficiency of PSII photochemistry (Fv/Fm) (b), in young and mature tomato leaves after 30 min of dark adaptation. Standard deviations (SDs) are shown by bars. Significant differences are shown by different lower-case letters (p < 0.05). Eight to ten plants were measured from each treatment (n = 8–10).
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Figure 2. The light energy partitioning at PSII. The effective quantum yield of PSII photochemistry (ΦPSII) (a); the quantum yield of regulated non-photochemical energy loss in PSII (ΦNPQ) (b); and the quantum yield of non-regulated energy loss in PSII (ΦNO) (c), in young and mature tomato leaves, evaluated at 426 μmol photons m−2 s−1 (low light intensity, LLI), and at 1000 μmol photons m−2 s−1 (high light intensity, HLI). Standard deviations (SDs) are shown by bars. Significant differences are shown by different lower-case letters (p < 0.05). Eight to ten plants were measured from each treatment (n = 8–10).
Figure 2. The light energy partitioning at PSII. The effective quantum yield of PSII photochemistry (ΦPSII) (a); the quantum yield of regulated non-photochemical energy loss in PSII (ΦNPQ) (b); and the quantum yield of non-regulated energy loss in PSII (ΦNO) (c), in young and mature tomato leaves, evaluated at 426 μmol photons m−2 s−1 (low light intensity, LLI), and at 1000 μmol photons m−2 s−1 (high light intensity, HLI). Standard deviations (SDs) are shown by bars. Significant differences are shown by different lower-case letters (p < 0.05). Eight to ten plants were measured from each treatment (n = 8–10).
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Figure 3. The fraction of open PSII reaction centers (RCs) (qp) (a), and the efficiency of the open PSII RCs (Fv’/Fm’) (b), in young and mature tomato leaves, evaluated at 426 μmol photons m−2 s−1 (low light intensity, LLI) and at 1000 μmol photons m−2 s−1 (high light intensity, HLI). Standard deviations (SDs) are shown by bars. Significant differences are shown by different lower-case letters (p < 0.05). Eight to ten plants were measured from each treatment (n = 8–10).
Figure 3. The fraction of open PSII reaction centers (RCs) (qp) (a), and the efficiency of the open PSII RCs (Fv’/Fm’) (b), in young and mature tomato leaves, evaluated at 426 μmol photons m−2 s−1 (low light intensity, LLI) and at 1000 μmol photons m−2 s−1 (high light intensity, HLI). Standard deviations (SDs) are shown by bars. Significant differences are shown by different lower-case letters (p < 0.05). Eight to ten plants were measured from each treatment (n = 8–10).
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Figure 4. The electron transport rate (ETR) (a), and the non-photochemical quenching (NPQ) (b), in young and mature tomato leaves, evaluated at 426 μmol photons m−2 s−1 (low light intensity, LLI) and at 1000 μmol photons m−2 s−1 (high light intensity, HLI). Standard deviations (SDs) are shown by bars. Significant differences are shown by different lower-case letters (p < 0.05). Eight to ten plants were measured from each treatment (n = 8–10).
Figure 4. The electron transport rate (ETR) (a), and the non-photochemical quenching (NPQ) (b), in young and mature tomato leaves, evaluated at 426 μmol photons m−2 s−1 (low light intensity, LLI) and at 1000 μmol photons m−2 s−1 (high light intensity, HLI). Standard deviations (SDs) are shown by bars. Significant differences are shown by different lower-case letters (p < 0.05). Eight to ten plants were measured from each treatment (n = 8–10).
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Figure 5. The excess excitation energy at PSII (EXC) (a) and the excitation pressure at PSII (1 − qL) (b) in young and mature tomato leaves, evaluated at 426 μmol photons m−2 s−1 (low light intensity, LLI) and at 1000 μmol photons m−2 s−1 (high light intensity, HLI). Standard deviations (SDs) are shown by bars. Significant differences are shown by different lower-case letters (p < 0.05). Eight to ten plants were measured from each treatment (n = 8–10).
Figure 5. The excess excitation energy at PSII (EXC) (a) and the excitation pressure at PSII (1 − qL) (b) in young and mature tomato leaves, evaluated at 426 μmol photons m−2 s−1 (low light intensity, LLI) and at 1000 μmol photons m−2 s−1 (high light intensity, HLI). Standard deviations (SDs) are shown by bars. Significant differences are shown by different lower-case letters (p < 0.05). Eight to ten plants were measured from each treatment (n = 8–10).
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Figure 6. Representative whole-leaf pseudocolor-coded pictures of the parameters Fv/Fm captured in young and mature dark-adapted tomato leaves, and of ΦPSII, ΦNPQ, and qp, captured at 426 μmol photons m−2 s−1 and at 1000 μmol photons m−2 s−1, in young and mature tomato leaves. The average whole-leaf value for each parameter is shown for each leaf. An asterisk indicates a significant difference at p < 0.05. At the bottom of the figure, the color code indicates the corresponding color values.
Figure 6. Representative whole-leaf pseudocolor-coded pictures of the parameters Fv/Fm captured in young and mature dark-adapted tomato leaves, and of ΦPSII, ΦNPQ, and qp, captured at 426 μmol photons m−2 s−1 and at 1000 μmol photons m−2 s−1, in young and mature tomato leaves. The average whole-leaf value for each parameter is shown for each leaf. An asterisk indicates a significant difference at p < 0.05. At the bottom of the figure, the color code indicates the corresponding color values.
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Moustaka, J.; Sperdouli, I.; Moustakas, M. Light Energy Use Efficiency in Photosystem II of Tomato Is Related to Leaf Age and Light Intensity. Crops 2024, 4, 623-635. https://doi.org/10.3390/crops4040043

AMA Style

Moustaka J, Sperdouli I, Moustakas M. Light Energy Use Efficiency in Photosystem II of Tomato Is Related to Leaf Age and Light Intensity. Crops. 2024; 4(4):623-635. https://doi.org/10.3390/crops4040043

Chicago/Turabian Style

Moustaka, Julietta, Ilektra Sperdouli, and Michael Moustakas. 2024. "Light Energy Use Efficiency in Photosystem II of Tomato Is Related to Leaf Age and Light Intensity" Crops 4, no. 4: 623-635. https://doi.org/10.3390/crops4040043

APA Style

Moustaka, J., Sperdouli, I., & Moustakas, M. (2024). Light Energy Use Efficiency in Photosystem II of Tomato Is Related to Leaf Age and Light Intensity. Crops, 4(4), 623-635. https://doi.org/10.3390/crops4040043

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