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The xanthophyll cycle and the water-water cycle had different functional significance in chilling-sensitive sweet pepper upon exposure to chilling temperature (4 °C) under low irradiance (100 µmol m−2 s−1) for 6 h. During chilling stress, effects of non-photochemical quenching (NPQ) on photosystem 2 (PS2) in dithiothreitol (DTT) fed leaves remained distinguishable from that of the water-water cycle in diethyldithiocarbamate (DDTC) fed leaves. In DTT-fed leaves, NPQ decreased greatly accompanied by visible inhibition of the de-epoxidized ratio of the xanthophyll cycle, and maximum photochemical efficiency of PS2 (Fv/Fm) decreased markedly. Thus the xanthophyll cycle-dependent NPQ could protect PS2 through energy dissipation under chilling stress. However, NPQ had a slighter effect on photosystem 1 (PS1) in DTT-fed leaves than in DDTC-fed leaves, whereas effects of the water-water cycle on PS1 remained distinguishable from that of NPQ. Inhibiting superoxide dismutase (SOD) activity increased the accumulation of $$\operatorname{O} \overline {^-- } _2$$ , the oxidation level of P700 (P700+) decreased markedly relative to the control and DTT-fed leaves. Both Fv/Fm and NPQ changed little in DDTC-fed leaves accompanied by little change of (A+Z)/(V+A+Z). This is the active oxygen species inducing PS1 photoinhibition in sweet pepper. The water-water cycle can be interrupted easily at chilling temperature. We propose that during chilling stress under low irradiance, the xanthophyll cycle-dependent NPQ has the main function to protect PS2, whereas the water-water cycle is not only the pathway to dissipate energy but also the dominant factor causing PS1 chilling-sensitivity in sweet pepper.

Chlorophyll fluorescence kinetics was used to investigate the effect of 1,4-dithiothreitol (DTT) on the distribution of excitation energy between photosystem 1 (PS1) and photosystem 2 (PS2) in soybean leaves under high irradiance (HI). The maximum PS2 quantum yield (Fv/Fm) was hardly affected by the presence of DTT, however, photon-saturated photosynthesis was depressed distinctly. Photochemical efficiency of open PS2 reaction centres during irradiation (Fv′/Fm′) was enhanced by about 30–40 % by DTT treatment, whereas photochemical quenching (qP) was depressed by about 40 % under HI. DTT treatment caused a 30 % decrease in allocation of excitation energy to PS1 under HI and a 20 % increase to PS2. An obvious shift in the balance of excitation energy distribution between photosystems was observed in DTT-treated leaves. Though high excitation pressure (1 - qP) resulted from DTT treatment, non-photochemical quenching (qN) was lower. DTT completely inhibited the formation of zeaxanthin and also distinctly depressed the state transition (qT). The shift in the balance of excitation distribution between the two photosystems induced by DTT was mainly due to the enhancement of excitation energy capture by PS2 antenna and the inhibition of state transition. It might be the shift in the balance between the two photosystems that mainly induced the depression of photosynthesis. Thus, to keep high utilization efficiency of absorbed photon energy, it is necessary to maintain the balance of excitation distribution between PS2 and PS1.

The effects of chilling treatment (4 °C) under low irradiance, LI (100 μmol m−2 s−1) and in the dark on subsequent recovery of photosynthesis in chilling-sensitive sweet pepper leaves were investigated by comparing the ratio of quantum yields of photosystem (PS) 2 and CO2 assimilation, ΦPS2/ΦCO2, measured in normal air (21 % O2, NA) and low O2-air (2% O2, LOA), and by analyzing chlorophyll (Chl) a fluorescence parameters. Chilling treatment in the dark had little effect on Fv/Fm and ΦPS2/ΦCO2, but it caused the decrease of net photosynthetic rate (PN) under saturating irradiance after 6-h chilling treatment, indicating that short-term chilling alone did not induce PS2 photoinhibition. Furthermore, photorespiration and Mehler reaction also did not obviously change during subsequent recovery after chilling stress in the dark. During chilling treatment under LI, there were obvious changes in Fv/Fm and ΦPS2/ΦCO2, determined in NA or LOA. Fv/Fm could recover fully in 4 h at 25 °C, and ΦPS2/ΦCO2 increased at the end of the treatment, as determined in both NA and LOA. During subsequent recovery, ΦPS2/ΦCO2 in LOA decreased faster than in NA. Thus the Mehler reaction might play an important role during chilling treatment under LI, and photorespiration was an important process during the subsequent recovery. The recovery of PN under saturating irradiance determined in NA and LOA took about 50 h, implying that there were some factors besides CO2 assimilation limiting the recovery of photosynthesis. From the progress of reduced P700 and the increase of the Mehler reaction during chilling under LI we propose that active oxygen species were the factors inducing PS1 photoinhibition, which prevented the recovery of photosynthesis in optimal conditions because of the slow recovery of the oxidizable P700.

The photoprotection of energy dissipation and water-water cycle were investigated by comparing chilling sensitivity of photosystems 2 (PS2) and 1 (PS1) in two chilling-sensitive plants, cucumber and sweet pepper, upon exposure to 4 °C under low irradiance (100 μmol m−2 s−1) for 6 h. During chilling stress, the maximum photochemical efficiency of PS2 (Fv/Fm) decreased only slightly in both plants, but the oxidisable P700 decreased markedly, which indicated that PS1 was more sensitive to chilling treatment under low irradiance than PS2. Sweet pepper leaves had lower Fv/Fm, higher non-photochemical quenching (NPQ), and higher oxidisable P700 during chilling stress. Activity of superoxide dismutase (SOD) and ascorbate peroxidase (APX) in cucumber leaves was higher, but APX activity decreased apparently compared to that at room temperature. The productions of active oxygen species (H2O2, O2−) increased in both plants, faster in cucumber leaves than in sweet pepper leaves. In sweet pepper leaves, a stronger de-epoxidation of the xanthophyll cycle pigments, a higher NPQ could act as a major protective mechanism to reduce the formation of active oxygen species during stress. Thus sensitivity of both plants to chilling under low irradiance was dominated by the protective mechanisms between PS1 and PS2, especially the energy dissipation and the water-water cycle.

Infiltration of methyl viologen (MV, source of O2−) and Na-diethyldithiocarbamate (DDC, inhibitor of SOD) into wheat leaves resulted in the accumulation of active oxygen species and photo-oxidative damage to photosynthetic apparatus under both moderate and high irradiance. Exogenous antioxidants, ascorbate (ASA) and mannitol, scavenged active oxygen efficiently, protected the photosynthetic system from MV and DDC induced oxidative damage, and maintained high Fv/Fm [maximal photochemical efficiency of photosystem 2 (PS2) while all PS2 reaction centres are open], Fm/F0 (another expression for the maximal photochemical efficiency of PS2), ΦPS2 (actual quantum yield of PS2 under actinic irradiation), qP (photochemical quenching coefficient), PN (net photosynthetic rate), and lowered qNP (non-photochemical quenching coefficient) of the leaves kept under high irradiance and oxidative stress. Phenolic compounds used in these experiments, catechol (Cat), resorcinol (Res), and tannic acid (Tan), had similar anti-oxidative activity and protective effect on photosynthetic apparatus as ASA and mannitol. The anti-oxidative activity and the protective effect of phenolic compounds increased with increase in their concentration from 100 to 300 g m−3. The number and the position of hydroxyl group in phenolic molecules seemed to influence their antioxidative activity.

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

Two cultivars of Capsicum annuum L. were acclimated for 5 d at sub-optimal temperature (14 °C) and irradiance of 250 µmol m−2 s−1. This cold-hardening resulted in some reduction in the extent of photoinhibition during an 8 h exposure to high irradiance at 4 °C. Obvious differences were observed between non-hardened leaves (NHL) and cold-hardened leaves (CHL) in the recovery under low irradiance at room temperature. The CHL of both cultivars recovered faster than NHL, especially during the initial fast phase of recovery. Compared with NHL, the total content of carotenoids (Cars), based on chlorophyll, Chl (a+b), and the proportions of xanthophyll cycle pigments referred to total Cars increased in CHL, mainly due to an increase of violaxanthin (V) + antheraxanthin (A) + zeaxanthin (Z) content per mol Chl (a+b). Faster development and a higher non-photochemical quenching (NPQ) of Chl fluorescence, related to a stronger deepoxidation of the larger xanthophyll cycle pool in NHL, could act as a major defence mechanism to reduce the formation of reactive oxygen species during severe chilling. This is suggested by higher content of Z or Z+A in photoinhibition as well as by its rapid decline during the initial fast phase of recovery. In contrast to the chilling-sensitive cv. 0004, the chilling-tolerant cv. 1141 did more easily acclimate its photosynthetic apparatus to low temperatures.

Pigment contents of chloroplasts and net photosynthetic rate were dramatically reduced in maize leaves suffering from iron deficiency. However, the reduction in photosynthesis was probably not caused by decreased contents of chlorophylls and carotenoids and by photon absorption; the primary limiting factor for photosynthesis may rather be the decrease of electron transport activity in photosystem 1. Iron-deficient leaves suffered serious acceptor-side photoinhibition, and more than 60 % of absorbed photons were dissipated, while less than 40 % was used in photochemical reaction. Thermal energy dissipation depending on xanthophyll cycle and D1 protein turnover was enhanced when acceptor-side photoinhibition occurred in iron-deficient maize leaves.

Changes in the activities of enzymes involved in scavenging active oxygen species were followed after exposing bean seedling leaves (Phaseolus vulgaris L.) to various cross stresses of irradiance and temperature. The activities of superoxide dismutase (SOD, EC 1.15.1.1) and ascorbate peroxidase (AsAPOD, EC 1.11.1.11) increased to different extent with prolonged irradiation of the leaves, and were stimulated by high temperature (HT). The activity of catalase (CAT, 1.11.1.6) decreased when exposed to strong irradiance (HI), and the decrease was further exacerbated when HI was combined with HT. CAT activity was more sensitive to HT than to HI. Ascorbate (AsA) content slightly decreased and then increased during the treatment of HI, but decreased under the cross stress of HI and HT. On the contrary, glutathione (GSH) content increased all the time during various treatments of irradiance and temperature. The increase in the combined stress was even more pronounced. Irradiance is the major reason in triggering the operation of xanthophyll cycle, which was difficult to be started by HT. The antioxidant systems tended to be inactivated with prolonged exposure to the cross stress of HI and HT. The de-expoxidated state of xanthophyll cycle, however, was increasing all the time, which indicated that the zeaxanthin-dependent thermal dissipation was one major energy dissipation pathway during the cross stress of HI and HT.

Photosynthesis in iron-deficient soybean and maize leaves decreased drastically. The quantum yield of photosystem 2 (PS2) electron transport (ΦPS2), the efficiency of excitation energy capture by open PS2 reaction centres (Fv′/Fm′), and photochemical quenching coefficient (qP) under high irradiance were lowered significantly by iron deficiency, but non-photochemical quenching (NPQ) increased markedly. The analysis of the polyphasic rise of fluorescence transient showed that iron depletion induced a pronounced K step both in soybean and maize leaves. The maximal quantum yield of PS2 photochemistry (Φpo) decreased only slightly, however, the efficiency with which a trapped exciton can move an electron into the electron transport chain further than QA (Ψ0) and the quantum yield of electron transport beyond QA (ΨEo) in iron deficient leaves decreased more significantly compared with that in control. Thus not only the donor side but also the acceptor of PS2 was probably damaged in iron deficient soybean and maize leaves.