Chilling stress and chilling tolerance of sweet potato as sensed by chlorophyll fluorescence

We studied changes in the chlorophyll (Chl) fluorescence components in chilling-stressed sweet potato (Ipomoea batatas L. Lam) cv. Tainung 57 (TN57, chilling-tolerant) and cv. Tainung 66 (TN66, chilling-susceptible). Plants under 12-h photoperiod and 400 μmol m⁻² s⁻¹ irradiance at 24/20 °C (day/night) were treated by a 5-d chilling period at 7/7 °C. Compared to TN66, TN57 exhibited a significantly greater basic Chl fluorescence (F₀), maximum fluorescence (F_m), maximum fluorescence yield during actinic irradiation (F_m′ ), and the quantum efficiency of electron transport through photosystem 2, PS2 (Φ_PS2). Chilling stress resulted in decrease in the potential efficiency of PS2 (F_v/F_m), Φ_PS2, non-photochemical fluorescence quenching (NPQ), non-photochemical quenching (q_N), and the occurrence of chilling injury in TN66. Chilling increased the likelihood of photoinhibition, characterized by a decline in the Chl fluorescence of both cultivars, and photoinhibition during low temperature stress generally occurred more rapidly in TN66.     

Involvement of betacyanin in chilling-induced photoinhibition in leaves of <Emphasis Type="Italic">Suaeda salsa</Emphasis>

Seeds of Suaeda salsa were cultured in dark for 3 d and betacyanin accumulation in seedlings was promoted significantly. Then the seedlings with accumulated betacyanin (C+B) were transferred to 14/10 h light/dark and used for chilling treatment 15 d later. Photosystem 2 (PS2) photochemistry, D1 protein content, and xanthophyll cycle during the chilling-induced photoinhibition (exposed to 5 °C at a moderate photon flux density of 500 μmol m⁻² s⁻¹ for 3 h) and the subsequent restoration were compared between the C+B seedlings and the control (C) ones. The maximal efficiency of PS2 photochemistry (F_v/F_m), the efficiency of excitation energy capture by open PS2 centres (F_v′/F_m′), and the yield of PS2 electron transport (Φ_PS2) of the C+B and C leaves both decreased during photoinhibition. However, smaller decreases in F_v/F_m, F_v′/F_m′, and Φ_PS2 were observed in the C+B leaves than in C ones. At the same time, the deepoxidation state of xanthophyll cycle, indicated by (A+Z)/(V+A+Z) ratio, increased rapidly but the D1 protein content decreased considerably during the photoinhibition. The increase in rate of (A+Z)/(V+A+Z) was higher but the D1 protein turnover was slower in C+B than C leaves. After photoinhibition treatment, the plants were transferred to a dim irradiation (10 μmol m⁻² s⁻¹) at 25 °C for restoration. During restoration, the chlorophyll (Chl) fluorescence parameters, D1 protein content, and xanthophyll cycle components relaxed gradually, but the rate and level of restoration in the C+B leaves was greater than those in the C leaves. The addition of betacyanins to the thylakoid solution in vitro resulted in similar changes of F_v/F_m, D1 protein content, and (A+Z)/(V+A+Z) ratio during the chilling process. Therefore, betacyanin accumulation in S. salsa seedlings may result in higher resistance to photoinhibition, larger slowing down of D1 protein turnover, and enhancement of non-radiative energy dissipation associated with xanthophyll cycle, as well as in greater restoration after photoinhibition than in the control when subjected to chilling at moderate irradiance.     

Effects of different irradiances on the photosynthetic process during ex-vitro acclimation of Anoectochilus plantlets

Six months old in vitro-grown Anoectochilus formosanus plantlets were transferred to ex-vitro acclimation under low irradiance, LI [60 μmol(photon) m⁻² s⁻¹], intermediate irradiance, II [180 μmol(photon) m⁻² s⁻¹], and high irradiance, HI [300 μmol(photon) m⁻² s⁻¹] for 30 d. Imposition of II led to a significant increase of chlorophyll (Chl) b content, rates of net photosynthesis (P _N) and transpiration (E), stomatal conductance (g _s), electron transfer rate (ETR), quantum yield of electron transport from water through photosystem 2 (Φ_PS2), and activity of ribulose-1,5-bisphosphate carboxylase/ oxygenase (RuBPCO, EC 4.1.1.39). This indicates that Anoectochilus was better acclimated at II compared to LI treatment. On the other hand, HI acclimation led to a significant reduction of Chl a and b, P _N, E, g _s, photochemical quenching, dark-adapted quantum efficiency of open PS2 centres (F_v/F_m), probability of an absorbed photon reaching an open PS2 reaction centre (F_v′/F_m′), ETR, Φ_PS2, and energy efficiency of CO₂ fixation (Φ_CO2/Φ_PS2). This indicates that HI treatment considerably exceeded the photo-protective capacity and Anoectochilus suffered HI induced damage to the photosynthetic apparatus. Imposition of HI significantly increased the contents of antheraxanthin and zeaxanthin (ZEA), non-photochemical quenching, and conversion of violaxanthin to ZEA. Thus Anoectochilus modifies its system to dissipate excess excitation energy and to protect the photosynthetic machinery.     

Photosynthesis of lichen symbiotic alga Trebouxia erici as affected by irradiance and osmotic stress

The relation between oxygen evolution rate (OER) and quantum yield of photochemical reactions in photosystem 2 (Φ_PS2) was examined in lichen symbiotic alga Trebouxia erici Ahmadjian (strain UTEX 911) exposed to different irradiances and osmotic stress (2 M sucrose for 60 h). Linear relationship was found between OER and Φ_PS2 in control cell suspension within irradiance range of 0 – 500 μmol m⁻² s⁻¹. Under osmotic stress, OER and Φ_PS2 were significantly reduced. Relation between OER and Φ_PS2 was curvilinear due to strong osmotically-induced inhibition of OER at high irradiance. The highest used irradiance (500 μmol m⁻² s⁻¹) was photoinhibitory for osmotically-stressed T. erici because non-photochemical quenching (NPQ) increased substantially. Energy-dependent quenching represented major part of NPQ increase. Osmotic stress led also to the reduction of capacity of photochemical processes in PS 2 (F_V/F_M) and increase in F₀/F_M. These changes indicated negative effects of osmoticum on structure and function of photosynthetic apparatus.     

Cryoproective role of ribitol in Xanthoparmelia somloensis

Thalli of Xanthoparmelia somloensis with natural content of polyols (control) and polyol-free thalli (acetone-rinsed) were used to study ribitol effects at low temperatures. Thalli segments were cultivated in ribitol concentration of 32 or 50 mM for 168 h at temperatures +5, 0, and ?5 °C. The chlorophyll fluorescence parameters (potential yield of photochemical reactions in PS 2 (variable to maximum fluorescence ratio, F_v/F_m), effective quantum yield of photochemical reactions in PS 2 (Φ_PS2), and non-photochemical quenching (NPQ) were monitored in 24-h intervals using an imaging system. The effect of 32 mM ribitol on F_v/F_m and Φ_PS2 was apparent only at ?5 °C, however, the effect was seen throughout the whole exposure. Surprisingly, 50 mM ribitol concentration treatment led to a decrease in F_v/F_m and Φ_PS2 and to an increase in NPQ values at ?5 °C, while no change was observed at 0 °C and +5 °C. Acetone-rinsing caused decrease of F_v/F_m, Φ_PS2 and NPQ.     

Photosystem 2 photochemistry and pigment composition of a yellow mutant of rice (<Emphasis Type="Italic">Oryza sativa</Emphasis> L.) under different irradiances

A yellow leaf colouration mutant (named ycm) generated from rice T-DNA insertion lines was identified with less grana lamellae and low thylakoid membrane protein contents. At weak irradiance [50 μmol(photon) m⁻² s⁻¹], chlorophyll (Chl) contents of ycm were ≈20 % of those of WT and Chl a/b ratios were 3-fold that of wild type (WT). The leaf of ycm showed lower values in the actual photosystem 2 (PS2) efficiency (Φ_PS2), photochemical quenching (q_P), and the efficiency of excitation capture by open PS2 centres 1 (F_v′/F_m′) than those of WT, except no difference in the maximal efficiency of PS2 photochemistry (F_v/F_m). With progress in irradiance [100 and 200 μmol(photon) m⁻² s⁻¹], there was a change in the photosynthetic pigment stoichiometry. In ycm, the increase of total Chl contents and the decrease in Chl a/b ratio were observed. Φ_PS2, q_P, and F_v′/F_m′ of ycm increased gradually along with the increase of irradiance but still much less than in WT. The increase of xanthophyll ratio [(Z+A)/(V+A+Z)] associated with non-photochemical quenching (q_N) was found in ycm which suggested that ycm dissipated excess energy through the turnover of xanthophylls. No significant differences in pigment composition were observed in WT under various irradiances, except Chl a/b ratio that gradually decreased. Hence the ycm mutant developed much more tardily than WT, which was caused by low photon energy utilization independent of irradiance.     

Modifications in Photosynthetic Pigments and Chlorophyll Fluorescence in 20-Year-Old Pine Trees after a Four-Year Exposure to Carbon Dioxide and Temperature Elevation

Changes in pigment composition and chlorophyll (Chl) fluorescence parameters were studied in 20 year-old Scots pine (Pinus sylvestris L.) trees grown in environment-controlled chambers and subjected to ambient conditions (CON), doubled ambient CO₂ concentration (EC), elevated temperature (ambient +2−6 °C, ET), or a combination of EC and ET (ECT) for four years. EC did not significantly alter the optimal photochemical efficiency of photosystem 2 (PS2; F_v/F_m), or Chl a+b content during the main growth season (days 150–240) but it reduced F_v/F_m and the Chl a+b content and increased the ratio of total carotenoids to Chl a+b during the ‘off season’. By contrast, ET significantly enhanced the efficiency of PS2 in terms of increases in F_v/F_m and Chl a+b content throughout the year, but with more pronounced enhancement in the ‘off season’. The reduction in F_v/F_m during autumn could be associated with the CO₂-induced earlier yellowing of the leaves, whereas the temperature-stimulated increase in the photochemical efficiency of PS2 during the ‘off season’ could be attributed to the maintenance of a high sink capacity. The pigment and fluorescence responses in the case of ECT showed a similar pattern to that for ET, implying the importance of the temperature factor in future climate changes in the boreal zone. This revised version was published online in August 2006 with corrections to the Cover Date.     

The Susceptibility of Cucumber and Sweet Pepper to Chilling Under Low Irradiance is Related to Energy Dissipation and Water-Water Cycle

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 (F_v/F_m) 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 F_v/F_m, 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. This revised version was published online in August 2006 with corrections to the Cover Date.     

Chilling and freezing stress in live oaks (Quercus section Virentes): intra- and inter-specific variation in PS II sensitivity corresponds to latitude of origin

Sensitivity to cold and freezing differs between populations within two species of live oaks (Quercus section Virentes Nixon) corresponding to the climates from which they originate. Two populations of Quercus virginiana (originating from North Carolina and north central Florida) and two populations of the sister species, Q. oleoides, (originating from Belize and Costa Rica) were grown under controlled climate regimes simulating tropical and temperate conditions. Three experiments were conducted in order to test for differentiation in cold and freezing tolerance between the two species and between the two populations within each species. In the first experiment, divergences in response to cold were tested for by examining photosystem II (PS II) photosynthetic yield (ΔF/F _m′) and non-photochemical quenching (NPQ) of plants in both growing conditions after short-term exposure to three temperatures (6, 15 and 30°C) under moderate light (400 μmol m⁻² s⁻¹). Without cold acclimation (tropical treatment), the North Carolina population showed the highest photosynthetic yield in response to chilling temperatures (6°C). Both ecotypes of both species showed maximum ΔF/F _m′ and minimum NPQ at their daytime growth temperatures (30°C and 15°C for the tropical and temperate treatments, respectively). Under the temperate treatment where plants were allowed to acclimate to cold, the Q. virginiana populations showed greater NPQ under chilling temperatures than Q. oleoides populations, suggesting enhanced mechanisms of photoprotective energy dissipation in the more temperate species. In the second and third experiments, inter- and intra-specific differentiation in response to freezing was tested for by examining dark-adapted F _v/F _m before and after overnight freezing cycles. Without cold acclimation, the extent of post-freezing declines in F _v/F _m were dependent on the minimum freezing temperature (0, −2, −5 or −10°C) for both populations in both species. The most marked declines in F _v/F _m occurred after freezing at −10°C, measured 24 h after freezing. These declines were continuous and irreversible over the time period. The North Carolina population, however, which represents the northern range limit of Q. virginiana, showed significantly less decline in F _v/F _m than the north central Florida population, which in turn showed a lower decline in Fv/F _m than the two Q. oleoides populations from Belize and Costa Rica. In contrast, after exposure to three months of chilling temperatures (temperate treatment), the two Q. virginiana populations showed no decline in F _v/F _m after freezing at −10°C, while the two Q. oleoides populations showed declines in F _v/F _m reaching 0.2 and 0.1 for Costa Rica and Belize, respectively. Under warm growth conditions, the two species showed different F ₀ dynamics directly after freezing. The two Q. oleoides populations showed an initial rise in F ₀ 30 min after freezing, followed by a subsequent decrease, while the Q. virginiana populations showed a continuous decrease in F ₀ after freezing. The North Carolina population of Q. virginiana showed a tendency toward deciduousness in response to winter temperatures, dropping 58% of its leaves over the three month winter period compared to only 6% in the tropical treatment. In contrast, the Florida population dropped 38% of its leaves during winter. The two populations of the tropical Q. oleoides showed no change in leaf drop during the 3-months winter (10% and 12%) relative to their leaf drop over the same timecourse in the tropical treatment. These results indicate important ecotypic differences in sensitivity to freezing and cold stress between the two populations of Q. virginiana as well as between the two species, corresponding to their climates of origin.     

Low-temperature limitation of primary photosynthetic processes in Antarctic lichens <Emphasis Type="Italic">Umbilicaria antarctica</Emphasis> and <Emphasis Type="Italic">Xanthoria elegans</Emphasis>

Temperature response curves of chlorophyll a fluorescence parameters were used to assess minimum sub-zero temperature assuring functioning of photosynthetic photochemical processes in photosystem II (PS II) of Antarctic lichens. Umbilicaria Antarctica and Xanthoria elegans were measured within the temperature range from −20 to +10°C by a fluorometric imaging system. For potential (F _V/F _M) and actual (Φ _II) quantum yields of photochemical processes the minimum temperature was found to be between −10 and −20°C. Non-photochemical quenching (NPQ) of absorbed excitation energy increased with temperature drop reaching maximum NPQ at −15°C. Image analysis revealed intrathalline heterogeneity of chlorophyll a fluorescence parameters with temperature drop. Temperature response of Φ _II exhibited an S-curve with pronounced intrathalline differences in X. elegans. The same relation was linear with only limited intrathalline difference in U. antarctica. The results showed that Antarctic lichen species were well adapted to sub-zero temperatures and capable of performing primary photosynthesis at −15°C.     

Photosystem 2 is more tolerant to high temperature in apple (<Emphasis Type="Italic">Malus domestica</Emphasis> Borkh.) leaves than in fruit peel

Tolerance of photosystem 2 (PS2) to high temperature in apple (Malus domestica Borkh. cv. Cortland) leaves and peel was investigated by chlorophyll a fluorescence (OJIP) transient after exposure to 25 (control), 40, 42, 44, and 46 °C in the dark for 30 min. The positive L-step was more pronounced in a peel than in leaves when exposed to 44 °C. Heat-induced K-step became less pronounced in leaves than in peel when exposed to 42 °C or higher temperature. Leaves had negative L-and K-steps relative to the peel. The decrease of oxygen-evolving complex (OEC) by heat stress was higher in the peel than in the leaves. OJIP transient from the 46 °C treated peel could not reach the maximum fluorescence (F_m). The striking thermoeffect was the big decrease in the relative variable fluorescence at 30 ms (V_I), especially in the leaves. Compared with the peel, the leaves had less decreased maximum PS2 quantum efficiency (F_v/F_m), photochemical rate constant (K_P), F_m and performance index (PI) on absorption basis (PI_abs) and less increased minimum fluorescence (F₀) and non-photochemical rate constant (K_N), but more increased reduction of end acceptors at PS1 electron acceptor side per cross section (RE₀/CS₀) and per reaction center (RE₀/RC₀), quantum yield of electron transport from Q_A ⁻ to the end acceptors (ϕ _R0) and total PI (PI_abs,total) when exposed to 44 °C. In conclusion, PS2 is more thermally labile than PS1. The reduction of PS2 activity by heat stress primarily results from an inactivation of OEC. PS2 was more tolerant to high temperature in the leaves than in the peel.     

Dissipation of Excitation Energy During Development of Photosystem 2 Photochemistry in Helianthus annuus

Etiolated sunflower cotyledons developed in complete darkness and lacking photosystem (PS) 2 were exposed to continuous 200 µmol(photon) m^–2 s^–1 white light for 1, 3, 6, 12, and 18 h prior to evaluations of excitation-energy dissipation using modulated chlorophyll a fluorescence. Photochemical potential of PS2, measured as the dark-adapted quantum efficiency of PS2 (F_V(M)/F_M), and thermal dissipation from the antenna pigment-protein complex, measured as the Stern-Volmer non-photochemical quenching coefficient (NPQ), increased to 12 h of irradiation. Following 12 h of irradiation, thermal dissipation from the antennae pigment-protein complex decreased while the efficiency of excitation capture by PS2 centers (F_V/F_M) and light-adapted quantum efficiency of PS2 (_PS2) continued to increase to 18 h of irradiation. The fraction of the oxidized state of Q_A, measured by the photochemical quenching coefficient (q_P), remained near optimal and was not changed significantly by irradiation time. Hence during the development of maximum photochemical potential of PS2 in sunflower etioplasts, which initially lacked PS2, enhanced thermal dissipation helps limit excitation energy reaching PS2 centers. Changes of the magnitude of thermal dissipation help maintain an optimum fraction of the oxidized state of Q_A during the development of PS2 photochemistry.     

Changes of Donor and Acceptor Side in Photosystem 2 Complex Induced by Iron Deficiency in Attached Soybean and Maize Leaves

Photosynthesis in iron-deficient soybean and maize leaves decreased drastically. The quantum yield of photosystem 2 (PS2) electron transport (Φ_PS2), the efficiency of excitation energy capture by open PS2 reaction centres (F_v′/F_m′), and photochemical quenching coefficient (q_P) 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 (Φ_po) decreased only slightly, however, the efficiency with which a trapped exciton can move an electron into the electron transport chain further than Q_A (Ψ₀) and the quantum yield of electron transport beyond Q_A (Ψ_Eo) 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. This revised version was published online in August 2006 with corrections to the Cover Date.     

Up-regulation of cyclic electron flow and down-regulation of linear electron flow in antisense-<Emphasis Type="Italic">rca</Emphasis> mutant rice

To investigate how excess excitation energy is dissipated in a ribulose-1,5-bisphospate carboxylase/oxygenase activase antisense transgenic rice with net photosynthetic rate (P _N) half of that of wild type parent, we measured the response curve of P _N to intercellular CO₂ concentration (C _i), electron transport rate (ETR), quantum yield of open photosystem 2 (PS2) reaction centres under irradiation (F_v′/F_m′), efficiency of total PS2 centres (Φ_PS2), photochemical (q_P) and non-photochemical quenching (NPQ), post-irradiation transient increase in chlorophyll (Chl) fluorescence (PITICF), and P700⁺ re-reduction. Carboxylation efficiency dependence on C _i, ETR at saturation irradiance, and F_v′/F_m′, Φ_PS2, and q_P under the irradiation were significantly lower in the mutant. However, NPQ, energy-dependent quenching (q_E), PITICF, and P700⁺ re-reduction were significantly higher in the mutant. Hence the mutant down-regulates linear ETR and stimulates cyclic electron flow around PS1, which may generate the ΔpH to support NPQ and q_E for dissipation of excess excitation energy.     

Role of calcium ion in protection against heat and high irradiance stress-induced oxidative damage to photosynthesis of wheat leaves

Cross stress of heat and high irradiance (HI) resulted in the accumulation of active oxygen species and photo-oxidative damage to photosynthetic apparatus of wheat leaves during grain development. Pre-treatment with calcium ion protected the photosynthetic system from oxidative damage by reducing O^-. ₂ production, inhibiting lipid peroxidation, and retarding electrolyte leakage from cell. Therefore, high F_v/F_m [maximal photochemical efficiency of photosystem 2 (PS2) while all PS2 reaction centres are open], F_m/F₀ (another expression for the maximal photochemical efficiency of PS2), Φ_PS2 (actual quantum yield of PS2 under actinic irradiation), q_P (photochemical quenching coefficient), and P _N (net photosynthetic rate) were maintained, and lower q_NP (non-photochemical quenching coefficient) of the leaves was kept under heat and HI stress. EGTA (a chelant of calcium ion) and LaCl₃ (a blocker of Ca²⁺ channel in cytoplasmic membrane) had the opposite effect. Thus Ca ion may help protect the photosynthetic system of wheat leaves from oxidative damage induced by the cross stress of heat and HI.     

Down-regulation of photosystem 2 efficiency and spectral reflectance in mango leaves under very low irradiance and varied chilling treatments

In order to elucidate the effects of chilling-stress at night on photosystem 2 (PS2) efficiency under dim irradiance (DI), mango leaves were chilled to varied extent (8–3 °C) and for varied duration (0–12 h) in growth cabinets in the dark, and then exposed to DI (20 μmol m⁻² s⁻¹ PPFD) at each chilling-temperature for 1 h. Chilling in the dark had little effect on F_v/F_m of mango leaves. But both the extent and duration of chilling pre-treatments significantly affected F_v’/F_m’ when leaves were exposed to DI. This down-regulation of PS2 efficiency was closely related to xanthophyll de-epoxidation, assessed as photochemical reflectance index (PRI) and calculated from leaf spectral reflectance [(R₅₃₁ − R₅₇₀)/(R₅₃₁ + R₅₇₀)], and non-photochemical quenching (NPQ). The down-regulation of PS2 is a defence mechanism initiated at predawn in winter to alleviate the damage of PS2 by the sudden and strong irradiation at sunrise. Mango leaves, transferred suddenly from warm and dark room to DI and chilling showed a slight down-regulation of PS2 efficiency, in spite of an increased xanthophyll de-epoxidation. This might have been due to the unavailability of some cofactors required for NPQ.     

Influence of water stress on leaf photosynthetic characteristics in wheat cultivars differing in their susceptibility to drought

A gradual reduction in leaf water potential (Ψ_leaf), net photosynthetic rate (P _N), stomatal conductance, and transpiration rate was observed in two drought tolerant (C 306 and K 8027) and two susceptible (RW 893 and 899) genotypes subjected to water stress. The extent of reduction was lower in K 8027 and C 306 and higher in RW 893 and RW 899. Rewatering the plants after 5 d of stress restored P _N and other gas exchange traits in all four cultivars. Water stress had no significant effect on variable to maximum fluorescence ratio (F_v/F_m) indicating that water stress had no effect on primary photochemistry of photosystem 2 (PS2). However, water stress reduced the efficiency of excitation energy transfer (F′_v/F′_m) and the quantum yield of electron transport (Φ_PS2). The reduction was more pronounced in susceptible cultivars. Water stress had no significant effect on photochemical quenching, however, the non-photochemical quenching increased by water stress.     

The effect of additional red irradiation on the photosynthetic apparatus of <Emphasis Type="Italic">Pisum sativum</Emphasis>

Pisum sativum (L.) plants were grown under “white” luminescent lamps, W [45 μ mol(quantum) m⁻² s⁻¹] or under the same irradiation supplemented with narrow spectrum red light-emitting diodes (LEDs), RE [λ_max = 660 nm, Δλ = 20 nm, 40 μmol(quantum) m⁻² s⁻¹]. Significant differences in the chlorophyll (Chl) a fluorescence parameters, degree of State 1–State 2 transition, and the pigment-protein contents were found in plants grown under differing spectral composition. Addition of red LEDs to the “white light” resulted in higher effective quantum yield of photosystem 2 (PS2), i.e. F′_v/F′_m, linear electron transport (ϕ_PS2), photochemical quenching (q_P), and lower non-photochemical quenching (q_N as well as NPQ). The RE plants were characterised by higher degree State 1–State 2 transition, i.e. they were more effective in radiant energy utilisation. Judging from the data of “green” electrophoresis of Chl containing pigment-protein complexes of plants grown under various irradiation qualities, the percentage of Chl in photosystem 2 (PS2) reaction centre complexes in RE plants was higher and there was no difference in the total Chl bound with Chl-proteins of light-harvesting complexes (LHC2). Because the ratio between oligomeric and monomeric LHC2 forms was higher in RE plants, we suggest higher LHC2 stability in these ones.     

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

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

Protective Effects of Exogenous Antioxidants and Phenolic Compounds on Photosynthesis of Wheat Leaves under High Irradiance and Oxidative Stress

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 F_v/F_m [maximal photochemical efficiency of photosystem 2 (PS2) while all PS2 reaction centres are open], F_m/F₀ (another expression for the maximal photochemical efficiency of PS2), _PS2 (actual quantum yield of PS2 under actinic irradiation), q_P (photochemical quenching coefficient), P _N (net photosynthetic rate), and lowered q_NP (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.