PHOTOINHIBITION OF MECHANICALLY STIMULABLE BIOLUMINESCENCE IN THE AUTOTROPHIC DINOFLAGELLATE CERATIUM FUSUS (PYRROPHYTA)1

Ceratium fusus (Ehrenb.) Dujardin was exposed to light of different wavelengths and photon flux densities (PFDs) to examine their effects on mechanically stimulable bioluminescence (MSL). Photoinhibition of MSL was proportional to the logarithm of PFD. Exposure to I μmol photons·m^?2s^?1 of broadband blue light (ca. 400–500 nm) produced near-complete photoinhibition (≥90% reduction in MSL) with a threshold at ca. 0.01 μmol photons·m^?2·s^?1. The threshold of photoinhibition was ca. an order of magnitude greater for both broadband green (ca. 500–580 nm) and red light (ca. 660–700 nm). Exposure to narrow spectral bands (ca. 10 nm half bandwidth) from 400 and 700 nm at a PFD of 0.1 μmol photons·m^?2·s^?1 produced a maximal response of photoinhibition in the blue wavelengths (peak ca. 490 nm). A photoinhibition response (≥ 10%) in the green (ca. 500–540 nm) and red wavelengths (ca. 680 nm) occurred only at higher PFDs (1 and 10 μmol photons·m^?2·s^?1). The spectral response is similar to that reported for Gonyaulax polyedra Stein and Pyrocystis lunula Schütt and unlike that of Alexandrium tamarense (Lebour) Balech et Tangen. The dinoflagellate's own bioluminescence is two orders of magnitude too low to result in self-photoinhibition. The quantitative relationships developed in the laboratory predict photoinhibition of bioluminescence in populations of C. fusus in the North Atlantic Ocean.     

Enhancement of susceptivity to photoinhibition and photooxidation in rice chlorophyll <Emphasis Type="Italic">b</Emphasis>-less mutants

Two rice chlorophyll (Chl) b-less mutants (VG28-1, VG30-5) and the respective wild type (WT) plant (cv. Zhonghua No. 11) were analyzed for the changes in Chl fluorescence parameters, xanthophyll cycle pool, and its de-epoxidation state under exposure to strong irradiance, SI (1 700 μmol m⁻² s⁻¹). We also examined alterations in the chloroplast ultrastructure of the mutants induced by methyl viologen (MV) photooxidation. During HI (0–3.5 h), the photoinactivation of photosystem 2 (PS2) appeared earlier and more severely in Chl b-less mutants than in the WT. The decreases in maximal photochemical efficiency of PS2 in the dark (F_v/F_m), quantum efficiency of PS2 electron transport (Φ_PS2), photochemical quenching (q_P), as well as rate of photochemistry (P_rate), and the increases in de-epoxidation state (DES) and rate of thermal dissipation of excitation energy (D_rate) were significantly greater in Chl b-mutants compared with the WT plant. A relatively larger xanthophyll pool and 78–83 % conversion of violaxanthin into antheraxanthin and zeaxanthin in the mutants after 3.5 h of HI was accompanied with a high ratio of inactive/total PS2 (0.55–0.73) and high 1–q_P (0.57–0.68) which showed that the activities of the xanthophyll cycle were probably insufficient to protect the photosynthetic apparatus against photoinhibition. No apparent difference of chloroplast ultrastructure was found between Chl b-less mutants and WT plants grown under low, LI (180 μmol m⁻² s⁻¹) and high, HI (700 μmol m⁻² s⁻¹) irradiance. However, swollen chloroplasts and slight dilation of thylakoids occurred in both mutants and the WT grown under LI followed by MV treatment. These typical symptoms of photooxidative damage were aggravated as plants were exposed to HI. Distorted and loose scattered thylakoids were observed in particular in the Chl b-less mutants. A greater extent of photoinhibition and photooxidation in these mutants indicated that the susceptibility to HI and oxidative stresses was enhanced in the photosynthetic apparatus without Chl b most likely as a consequence of a smaller antenna size.     

GROWTH,PHOTOSYNTHESIS AND MAINTENANCE METABOLIC COST IN THE DIATOM PHAEODACTYLUM TRICORNUTUM AT VERY LOW LIGHT LEVELS 1

The compensation point for growth of Phaeodactylum tricornutum Bohlin is less than 1 μmol. m^?2s^?1. Growth at low PFDs (<3.5 μmol. m^?2.s^?1) does not appear to reduce the maximum quantum efficiency of photosynthesis (ø_m) or to greatly inhibit the potential for light-saturated, carbon-specific photosynthesis (P_m^c). The value for ø_m in P. tricornutum is 0.10–0.12 mol O₂-mol photon^?1, independent of acclimation PFD between 0.75 and 200 μmol.m^?2.s^?1 in nutrient-sufficient cultures. P_m^c in cells of P. tricornutum acclimated to PFDs <3.5 μmol m^?2?s^?1 is approximately 50% of the highest value obtained in nutrient-sufficient cultures acclimated to growth-rate-saturating PFDs. In addition, growth at low PFDs does not severely restrict the ability of cells to respond to an increase in light level. Cultures acclimated to growth at lees than 1% of the light-saturated growth rate respond rapidly to a shift-up in PFD after a short initial lag period and achieve exponential growth rates of 1.0 d^?1 (65% of the light- and nutrient-saturated maximum growth rate) at both 40 and 200 μmol.m^?2.s^?1     

The analysis of determining factors and evaluation of tolerance to photoinhibition in wheat (<Emphasis Type="Italic">Triticum aestivum</Emphasis> L.)

Photoinhibition is a significant constraint for improvement of radiation-use efficiency and yield potential in cereal crops. In this work, attached fully expanded leaves of seedlings were used to assay the factors determining photoinhibition and for evaluation of tolerance to photoinhibition in wheat (Triticum aestivum L.). Our results showed that even 1 h under PPFD of 600 µmol(photon) m^?2 s^?1 could significantly reduce maximal quantum yield of PSII photochemistry (F_v/F_m) and performance index (PI) compared to low light [300 µmol(photon) m^?2 s^?1]. The decrease of F_v/F_m and PI was more noticeable with the increase of light intensity; irradiance higher than 800 µmol(photon) m^?2 s^?1 resulted in photoinhibition. Compared to 25°C, lower (20°C) or higher temperature (≥ 35°C) aggravated photoinhibition, while slightly high temperature (28°) alleviated photoinhibition. At 25°C, irradiance of 1,000 µmol(photon) m^–2 s^–1 for 1 h was enough to cause photoinhibition and a significant decrease of F_v/F_m, PI, trapped energy flux, electron transport flux, and density of reaction center as well as increase of dissipated energy flux per cross section were observed. In addition, seedlings at 21–32 days after planting showed a relatively stable phenotype, while the younger or older seedlings indicated an increased susceptibility to photoinhibition, especially in senescing leaves. Finally, six wheat varieties with relative tolerance to photoinhibition were identified from 22 Chinese winter wheat varieties by exposing attached leaves of the 25-d old seedlings for 1 h to 1,000 µmol(photon) m^–2 s^–1 at 25°C. Therefore, our work established a possible method for development of new wheat varieties with enhanced tolerance to photoinhibition.     

Photosynthesis and non-photochemical excitation quenching components of chlorophyll excitation in maize and field bean during chilling at different photon flux density

The influence of chilling (8 °C, 5 d) at two photon flux densities [PFD, L = 200 and H = 500 μmol(photon) m⁻² s⁻¹] on the gas exchange and chlorophyll fluorescence was investigated in chilling-tolerant and chilling-sensitive maize hybrids (Zea mays L., K383×K130, K185×K217) and one cultivar of field bean (Vicia faba L. minor, cv. Nadwiślański). The net photosynthetic rate (P _N) for the both studied plant species was inhibited at 8 °C. P _N of both maize hybrids additionally decreased during chilling. Changes in the quantum efficiency of PS2 electron transport (Φ_PS2) as a response to chilling and PFD were similar to P _N. Measurements of Φ_PS2/Φ_CO2 ratio showed that in field bean seedlings strong alternative photochemical sinks of energy did not appear during chilling. However, the high increment in Φ_PS2/Φ_CO2 for maize hybrids can indicate reactions associated with chill damage generation. At 8 °C the non-photochemical quenching (NPQ) increased in all plants with chilling duration and PFD. The appearance of protective (q_I,p) and damage (q_I,d) components of q_I and a decrease in q_E (energy dependent quenching) took place. NPQ components of field bean and maize hybrids differed from each other. The amount of protective NPQ (q_E + q_I,p) components as part of total NPQ was higher in field bean than in maize hybrids at both PFD. On 5^th day of chilling, the sum of q_E and q_I,p was 26.7 % of NPQ in tolerant maize hybrids and 17.6 % of NPQ in the sensitive one (averages for both PFD). The increased PFD inhibited the ability of all plants to perform protective dissipation of absorbed energy. The understanding of the genotypic variation of NPQ components in maize may have implications for the future selection of plants with a high chilling tolerance.     

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.     

Genotypic variation within Zea mays for susceptibility to and rate of recovery from chill-induced photoinhibition of photosynthesis

Six genotypes of Zea mays L. were grown in pots inside a glasshouse at a mean temperature of 22±2°C and a minimum photosynthetic photon flux density (Q) during the daylight period of 400 μmol m^?2 s^?1. Chilling-dependent photoinhibition was induced by exposing plants to a temperature of 7°C and a Q of 1 000 μmol m^?2 s^?1 for 6 h. Recovery from photoinhibition was then followed at a temperature of 25°C and a Q of 200 μmol m^?2 s^?1. Leaf gas exchange and chlorophyll fluorescence were measured on attached leaves at room temperature prior to the photoinhibitory treatments and at 6 sampling intervals from 0 to 24 h during the recovery period. The relative water content (RWC) was also measured during the recovery period. The results showed a significant genotypic variation in the susceptibility to and rate of recovery from chilling-dependent photoinhibition of photosynthesis in Zea mays seedlings. The Highland Pool 1a from highland sites in Mexico was the least susceptible to chill-induced photoinhibition, but had the slowest rate of recovery. The hybrid variety LG11 showed the highest rate of recovery, whilst the inbred line ZPF307 was the most susceptible to chill-induced photoinhibition. Susceptibility to photoinhibition and subsequent recovery were at least partially independent, suggesting that selection for improved genotypes will require independent selection for both tolerance and capacity for recovery. Although chlorophyll fluorescence provided a more rapid method of assessing the occurrence of photoinhibition, it was not as effective as direct gas-exchange measurements of the maximum quantum yield of photosynthesis (φ) in separating genotypes with respect to their susceptibility to photoinhibition, especially in the most vulnerable genotypes such as ZPF307. Water stress induced by chilling and high Q treatments appeared to impair the recovery processes. Decreases in stomatal conductance (g_s) produce a significant decrease in intercellular CO₂ concentration (C_i), although this decrease was never so extreme that it limited photosynthetic rates at the light intensities used to determine φ. Nevertheless, closure of stomata in patches, producing local restriction of CO₂ supply, would explain the poor correlation between chlorophyll fluorescence and quantum yield measurements in some genotypes immediately after photoinhibitory treatments.     

Recovery of photosynthesis and phtosystem II fluorescence in Chlamydomonas reinhardtii after exposure to three levels of high light

Recovery from 60 min of photoinhibitory treatment at photosynthetic photon flux densities of 500, 1400 and 2200 μMmol m^?2 s^? was followed in cells of the green alga Chlamydomonas reinhardtii grown at 125 μMmol m^?2 s^?1. These light treatments represent photoregulation, moderate photoinhibition and strong photoinhibition, respectively. Treatment in photoregulatory light resulted in an increased maximal rate of oxygen evolution (P_max) and an increased quantum yield (Φ), but a 15% decrease in F_v/F_M. Treatment at moderately photoinhibitory light resulted in a 30% decrease in F_v/F_M and an approximately equal decrease in Φ. Recovery in dim light restored F_v/F_M within 15 and 45 min after high light treatment at 500 and 1400 μMmol m^?2 s^?1, respectively. Convexity (Θ), a measure of the extent of co-limitation between PS II turnover and whole-chain electron transport, and Φ approached, but did not reach the control level during recovery after exposure to 1400 μMmol m^?2 s^?1, whereas P_max increased above the control. Treatment at 2200 μMmol m^?2 s^?1 resulted in a strong reduction of the modeled parameters Φ, Θ and P_max. Subsequent recovery was initially rapid but the rate decreased, and a complete recovery was not reached within 120 min. Based on the results, it is hypothesized that exposure to high light results in two phenomena. The first, expressed at all three light intensities, involves redistribution within the different aspects of PS II heterogeneity rather than a photoinhibitory destruction of PS II reaction centers. The second, most strongly expressed at 2200 μmol m^?2 s^?1, is a physical damage to PS II shown as an almost total loss of PS II_α and PS II Q_B-reducing centers. Thus recovery displayed two phase, the first was rapid and the only visible phase in algae exposed to 500 and 1400 μmol m^?2 s^?1. The second phase was slow and visible only in the later part of recovery in cells exposed to 2200 μmol m^?2 s^?1.     

Relationship Between Photosystem 2 Electron Transport and Photosynthetic CO2 Assimilation Responses to Irradiance in Young Apple Tree Leaves

The responses to irradiance of photosynthetic CO₂ assimilation and photosystem 2 (PS2) electron transport were simultaneously studied by gas exchange and chlorophyll (Chl) fluorescence measurement in two-year-old apple tree leaves (Malus pumila Mill. cv. Tengmu No.1/Malus hupehensis Rehd). Net photosynthetic rate (P _N) was saturated at photosynthetic photon flux density (PPFD) 600-1 100 (mol m^-2 s^-1, while the PS2 non-cyclic electron transport (P-rate) showed a maximum at PPFD 800 mol m^-2 s^-1. With PPFD increasing, either leaf potential photosynthetic CO₂ assimilation activity (F_d/F_s) and PS2 maximal photochemical activity (F_v/F_m) decreased or the ratio of the inactive PS2 reaction centres (RC) [(F_i – F_o)/(F_m – F_o)] and the slow relaxing non-photochemical Chl fluorescence quenching (q_s) increased from PPFD 1 200 mol m^-2 s^-1, but cyclic electron transport around photosystem 1 (RFp), irradiance induced PS2 RC closure [(F_s – F_o)/F_m – F_o)], and the fast and medium relaxing non-photochemical Chl fluorescence quenching (q_f and q_m) increased remarkably from PPFD 900 (mol m^-2 s^-1. Hence leaf photosynthesis of young apple leaves saturated at PPFD 800 mol m^-2 s^-1 and photoinhibition occurred above PPFD 900 mol m^-2 s^-1. During the photoinhibition at different irradiances, young apple tree leaves could dissipate excess photons mainly by energy quenching and state transition mechanisms at PPFD 900-1 100 mol m^-2 s^-1, but photosynthetic apparatus damage was unavoidable from PPFD 1 200 mol m^-2 s^-1. We propose that Chl fluorescence parameter P-rate is superior to the gas exchange parameter P _N and the Chl fluorescence parameter F_v/F_m as a definition of saturation irradiance and photoinhibition of plant leaves.     

Chlorophyll fluorescence and photoinhibition in a tropical rainforest understory plant

The data presented here deal with the effects of high-light exposure on the 77 K fluorescence characteristics of Elatostema repens. It is shown that the decrease of the variable fluorescence during the treatment is biphasic. The reactions responsible for the first phase of fluorescence quenching are saturated under 700 mol photon m^-2 s^-1 and insensitive to streptomycin, whereas those responsible for the second phase are not yet saturated under 700 mol photon m^-2 s^-1 and sensitive to streptomycin. It is concluded that only the second phase of fluorescence quenching is associated with photoinhibitory processes. Rate and amplitude of recovery from photoinhibition are maximum under very low light (3.5 mol photon m^-2 s^-1), and very small at a moderate light (160 mol photon m^-2 s^-1) which does not cause photoinhibition. It is concluded that recovery processes are inhibited during photoinhibition. It is suggested that they could be associated with damage occuring on the oxidizing side of PSII.Abbreviations F_o, F_v, F_m initial, variable and maximum fluorescence, respectively - PFD photon flux density - PS II photosystem II     

Photosystem 2 activity of <Emphasis Type="Italic">Citrus volkameriana</Emphasis> (L.) leaves as affected by Mn nutrition and irradiance

Citrus volkameriana (L.) plants were grown for 43 d in nutrient solutions containing 0, 2, 14, 98, or 686 μM Mn (Mn₀, Mn₂, Mn₁₄, Mn₉₈, and Mn₆₈₆, respectively). To adequately investigate the combined effects of Mn nutrition and irradiance on photosystem 2 (PS2) activity, irradiance response curves for electron transport rate (ETR), nonphotochemical quenching (q_N), photochemical quenching (q_P), and real photochemical efficiency of PS2 (Φ_PS2) were recorded under 10 different irradiances (66, 96, 136, 226, 336, 536, 811, 1 211, 1 911, and 3 111 μmol m⁻² s⁻¹, I₆₆ to I₃₁₁₁, respectively) generated with the PAM-2000 fluorometer. Leaf chlorophyll content was significantly lower under Mn excess (Mn₆₈₆) compared to Mn₀; its highest values were recorded in the treatments Mn₂-Mn₉₈. However, ETR and Φ_PS2 values were significantly lower under Mn₀ compared to the other Mn treatments, when plants were exposed to irradiances ≥96 μmol m⁻² s⁻¹. Furthermore, Mn₀ plants had significantly higher values of q_N and lower values of q_P at irradiances ≤226 and ≥336 μmol m⁻² s⁻¹, respectively, than those grown under Mn₂-Mn₆₈₆. Irrespective of Mn treatment, the values of Φ_PS2 and q_N decreased, while those of q_P increased progressively by increasing irradiance from I₁₃₆ to I₃₁₁₁. Finally, Mn₂-Mn₉₈ plants were less sensitive to photoinhibition of photosynthesis (≥811 μmol m⁻² s⁻¹) than the Mn₆₈₆ (≥536 μmol m⁻² s⁻¹) and Mn₀ (≥336 μmol m⁻² s⁻¹) ones.     

Chlorophyll fluorescence in micropropagated <Emphasis Type="Italic">Rhododendron ponticum</Emphasis> subsp. <Emphasis Type="Italic">baeticum</Emphasis> plants in response to different irradiances

The aim of this study was to investigate acclimation of micropropagated plants of Rhododendron ponticum subsp. baeticum to different irradiances and recovery after exposure to high irradiance. Plants grown under high (HL) or intermediate (IL) irradiances displayed higher values of maximum electron transport rate (ETR_max) and light saturation coefficient (E_k) than plants grown under low irradiance (LL). The capacity of tolerance to photoinhibition (as assessed by the response of photochemical quenching, q_p) varied as follows: HL > IL > LL. Thermal energy dissipation (q_N) was also affected by growth irradiance, with higher saturating values being observed in HL plants. Light-response curves suggested a gradual replacement of q_p by q_N with increasing irradiance. Following exposure to irradiance higher than 1500 μmol m⁻² s⁻¹, a prolonged reduction of the maximal photochemical efficiency of PS 2 (F_v/F_m) was observed in LL plants, indicating the occurrence of chronic photoinhibition. In contrary, the decrease in F_v/F_m was quickly reverted in HL plants, pointing to a reversible photoinhibition.     

Temperature response and photoinhibition investigated by chlorophyll fluorescence measurements for four distinct species of dipterocarp trees

The effects of strong light in combination with elevated temperatures on the photosynthetic system were examined in 4 dipterocarp tree species with ecologically different habitats. The 4 dipterocarp tree species were: Shorea platyclados originated from upper dipterocarp forests, Shorea parvifolia– lowland and hill dipterocarp forests, Shorea assamica– lowland dipterocarp forests, and Dipterocarpus oblongifolius– riparian fringes. S. platyclados and D. oblongifolius have higher growth and survival rates in open sites than S. parvifolia and S. assamica. Tolerance of high temperature among the species was assessed by determining the critical temperatures (T_c) at which the minimal fluorescence (F_o) began to rise sharply. This was measured by exposing plants to an increasing temperature of about 1°C min^?1. The intrinsic thermotolerance of the thylakoid membrane appears to be the highest for D. oblongifolius (T_c=46.4°C), intermediate for S. platyclados (45.7°C), and lowest for S. parvifolia and S. assamica (45.2 and 45.3°C, respectively). The temperature‐dependent efficiency of PSII electron transport (ΔF/F′_m), photochemical quenching (q_P), and the efficiency of light capture of open PSII (F′_v/F′_m) were measured at the photosynthetic steady state at least 10 min after the light exposure (180 μmol m^?2 s^?1 PFD). Stable temperature responses of ΔF/F′_m and q_P were observed in S. platyclados and D. oblongifolius, while those in S. parvifolia and S. assamica were more temperature‐dependent and severely affected at 45°C. Little difference was observed in temperature‐dependent F′_v/F′_m among species. Photoinhibitory light exposure (1600 μmol m^?2 s^?1 PFD) for 2 h at 40°C had little effect on the recovery kinetics from photoinhibition of S. platyclados and D. oblongifolius compared with those at 35°C. In contrast, the recovery from photoinhibition was retarded in S. parvifolia and S. assamica. These findings suggest that even at 40°C, a temperature below T_c, an exposure to strong light exacerbated photoinhibition in S. parvifolia and S. assamica corresponding to the closure of PSII reaction centers, as indicated by the decrease in q_P at this temperature. Thus, S. platyclados and D. oblongifolius, which occur at uplands and riparian fringes with frequent disturbances, are suggested to have higher photosynthetic tolerance to elevated temperatures contributing to a circumvention of photoinhibition.     

Chlorophyll fluorescence quenching and violaxanthin deepoxidation of FBPase antisense plants at low light intensities and low temperatures

Genetically modified potato (Solanum tuberosum L. cv. Desiree) and tobacco (Nicotiana tabacum cv. Samsun N.N.) plants were used to analyze the effects exerted by the chloroplastic (cp) fructose- 1,6-bisphosphatase (FBPase) on the regulation of light energy discrimination at the level of photosystem II. The cp-FBPase activity was progressively inhibited by an mRNA antisense to this FBPase. The chlorophyll fluorescence quenching parameters of these transgenic plants were compared to those of wild-type and transgenic plants that were acclimated to low temperatures. In particular various lines of the transgenic potato and tobacco plants were exposed to a temperature treatment of 10 and 20°C for 10 days. Light intensities were kept low to reduce photoinhibition so that we could analyze exclusively the effects of a modification in the carbon fixation cycle on the chlorophyll fluorescence quenching parameters. The photon flux densities (PFDs) employed at the level of the middle leaves of all plants were set to two different values of 10 μmol m^?2 s^?1 and 50 μmol m^?2 s^?1. Subsequent to this 10-day acclimation the chlorophyll-fluorescence parameters of all plants were measured. Photoinhibition as expressed by the F_y/F_m ratio was minor in plants subjected to a PFD of 10 μmol m^?2 s^?1. Higher photon fluence rates of 50 μmol m^?2 s^?1 at temperatures of 10°C gave rise to a significant reduction in the F_y/F_m ratios obtained from the transgenic plants which were characterized by a restriction in cp-FBPase capacity to 20% of normal activity. Furthermore, a progressive inhibition of the cp-FBPase activity induced an amplified nonphotochemical quenching of chlorophyll fluorescence with in the genetically manipulated species (except at 10°C and 50 μmol m^?2 s^?1). The increase in nonphotochemical quenching depended upon light and temperature. Photochemical quenching of light quanta within the antisense plants declined relative to that in the wild type. To further characterize the mechanisms producing higher levels of nonphotochemical fluorescence quenching. we analyzed several of the xanthophyll cycle pigments. The deepoxidation state of the xanthophyll cycle pigments in potato plants increased with attenuating FBPase activities under all conditions. For tobacco plants, this elevation of the deepoxidation state was only observed at a PFD of 50 μmol m^?2 s^?1.     

THE INTERACTION BETWEEN INORGANIC CARBON ACQUISITION AND LIGHT SUPPLY IN PALMARIA PALMATA (RHODOPHYTA)1

The dependence of the carbon concentrating mechanism of Palmaria palmata (L.) Kuntze on the growth light level was examined 1) to determine whether or not there is a threshold photon flux density (PFD) at which the inorganic carbon uptake mechanism can operate and 2) to attempt to quantify the relative energetic costs of acclimation to the two different limiting factors, PFD and dissolved inorganic carbon (DIC) concentration. Plants were grown at six PFDs: 5, 25, 50, 75, 95, and 125 μmol photons. m^?2.s^?1. Growth rates increased with increasing PFD from 5 to 50 μmol photons. m^?2. s^?1 and were light-saturated at 75, 95, and 125 μmol photons. m^?2. s^?1 Values of δ¹³C increased continuously with increasing growth PFD and did not saturate over the range of light levels tested. Time-resolved fluorescence characteristics indicated a progressive photoacclimation below 50 μmol photons. m^?2. s^?1. Analysis of chlorophyll fluorescence induction showed three levels of light use efficirncy associated with growth at 5 or 25, 50, and >75 μmol photons. m^?2. s^?1. The light-haruesting efficiency was inversely proportional to the effectiveness of DIC acquisition in plants grown at the six PFDs. These data were interpreted to indicate that there is a physiological tradeoff between photosynthetic efficiency and bicarbonate use in this species.     

Photosynthetic performance and resistance to photoinhibition of Zea mays L. leaves grown at sub-optimal temperature

The performance of the photosynthetic apparatus was examined in the third leaves of Zea mays L. seedlings grown at near-optimal (25 °C) or at suboptimal (15 °C) temperature by measuring chlorophyll (ChI) a fluorescence parameters and oxygen evolution in different temperature and light conditions. In leaf tissue grown at 25 and 15 °C, the quantum yield of PSII electron transport (ψ_PSII) and the rate of O₂ evolution decreased with decreasing temperature (from 25 to 4 °C) at a photon flux density of 125 μmol m^?2 s^?1. In leaves grown at 25 °C, the decrease of ψ_PSII correlated with a decrease of photochemical ChI fluorescence quenching (q_p), whereas in leaves crown at 15 °C q_p was largely insensitive to the temperature decrease. Compared with leaves grown at 25 °C, leaves grown at 15 °C were also able to maintain a higher fraction of oxidized to reduced Q_A (greater q_p) at high photon flux densities (up to 2000 μmol m^?2 s^?1), particularly when the measurements were performed at high temperature (25 °C). With decreasing temperature and/or increasing light intensity, leaves grown at 15 °C exhibited a substantial quenching of the dark level of fluorescence F₀ (q₀) whereas this type of quenching was virtually absent in leaves grown at 25 °C. Furthermore, leaves grown at 15 °C were able to recover faster from photo inhibition of photosynthesis after a photoinhibitory treatment (1200 μmol m^?2 s^?1 at 25, 15 or 6 °C for 8 h) than leaves grown at 25 °C. The results suggest that, in spite of having a low photosynthetic capacity, Z. mays leaves grown at sub optimal temperature possess efficient mechanisms of energy dissipation which enable them to cope better with photoinhibition than leaves grown at near-optimal temperature. It is suggested that the resistance of Z. mays leaves grown at 15 °C to photoinhibition is related to the higher content of carotenoids of the xanthophyll cycle (violaxanthin + antheraxanthin + zeaxanthin) measured in these leaves than in leaves grown at 25 °C.     

The impact of ozone fumigation and fertilization on chlorophyll fluorescence of birch leaves (Betula pendula)

 The impact of ozone fumigation on chlorophyll a fluorescence parameters and chlorophyll content of birch trees grown at high and low fertilization were studied for 6-, 8-, and 12-week old leaves. Fluorescence parameters were measured with a portable fluorometer with its fibre optics tightly inserted in a gas exchange cuvette at light intensities from 0 to 220 μmol photons m⁻² s⁻¹. Ozone caused significant changes of primary photosynthetic reactions: a decrease of the quantum yield of photosystem II and an increase of non-photochemical quenching. In all leaves a biphasic light response of non-photochemical quenching was observed. Ozone fumigation shifted the onset of the second phase from a PFD of about 60 μmol m⁻² s⁻¹ to about 30 μmol m⁻² s⁻¹. While the fertilizer concentration had no influence on this character, high fertilization supply of plants partially reduced O₃-induced damage. The light responses of Ft, Fm′ and NPQ observed in birch leaves grown in O₃-free air indicate the existence of at least two different processes governing energy conversion of the photosynthetic apparatus at PS II in the range of PFD 0–200 μmol photons m⁻² s⁻¹. The first phase was attributed to a rather slowly relaxing type of non-photochemical quenching, which, at least at low PFD, is thought to be related to a state 1–2 transition. The further changes of the fluorescence parameters studied at higher PFD might be explained by an increase of energy-dependent quenching, connected with the energization of the thylakoid membrane and zeaxanthin synthesis. A major effect of ozone treatment was a lowering of PS II quantum yield. This reflects a reduction of PS II electron transport and corresponds to the reduction of CO₂-fixation observed in ozonated leaves. Received: 24 September 1996 / Accepted: 27 January 1999     

Effects of chilling and high temperatures on photosynthesis and chlorophyll fluorescence in leaves of watermelon seedlings

The effects of chilling (CT, day/night temperatures of 12/10 °C, an irradiance of 250 μmol m^?2 s^?1), chilling combined with a low irradiance (CL, 12/10 °C, 80 μmol m^?2 s^?1), and a high temperature (HT, 42/40 °C, 250 μmol m^?2 s^?1) on chlorophyll content, chlorophyll fluorescence, and gas exchange were studied in two watermelon cultivars, ZJ8424 and YS01, differing in their resistance. The chlorophyll content, net photosynthetic rate (P_N), stomatal conductance (g_s), and transpiration rate (E) decreased substantially, whereas the intercellular CO₂ concentration (c_i) increased when the two watermelon cultivars were grown under these stresses. The photosynthetic parameters showed greater changes at chilling than at the high temperature, and the CL caused a more pronounced inhibition in PN compared with the CT. After 2 d exposure to the CT, YS01 had higher P_N, g_s, and E, but a lower ci compared with ZJ8424. The maximum efficiency of photosystem (PS) II photochemistry (F_v/F_m), effective quantum yield of PS II photochemistry (Φ_PSII), photochemical quenching (qP), and electron transport rate (ETR) decreased under the CT and CL but showed only a slight drop under the HT. All these stresses significantly increased non-photochemical quenching (NPQ). The CT brought more damage to the photosynthetic apparatus of leaves compared with the CL. In addition, after returning to normal conditions (25/15 °C, 250 μmol m^?2 s^?1) for 3 d, the photosynthetic parameters recovered to pre-stress levels in HT treated seedlings but not in CT treated seedlings. In conclusion, the low irradiance could help to alleviate the extent of photoinhibition of PS II photochemistry caused by chilling and cv. ZJ8424 was more sensitive to the extreme temperatures than cv. YS01.     

Photosynthesis of Marine Macroalgae from Antarctica: Light and Temperature Requirements

The photosynthetic performance of macroalgae isolated in Antarctica was studied in the laboratory. Species investigated were the brown algae Himantothallus grandifolius, Desmarestia anceps, Ascoseira mirabilis, the red algae Palmaria decipiens, Iridaea cordata, Gigartina skottsbergii, and the green algae Enteromorpha bulbosa, Acrosiphonia arcta, Ulothrix subflaccida and U. implexa. Unialgal cultures of the brown and red algae were maintained at 0°C, the green algae were cultivated at 10°C. I_K values were between 18 and 53 μmol m^?2 s^?1 characteristic or low light adapted algae. Only the two Ulothrix species showed higher I_K values between 70 and 74 μmol m^?2 s^?1. Photosynthesis compensated dark respiration at very low photon fluence rates between 1.6 and 10.6 μmol m^?2 s^?1. Values of α were high: between 0.4 and 1.1 μmol O₂ g^?1 FW h^?1 (μmol m^?2 s^?1)^?1 in the brown and red algae and between 2.1 and 4.9 μmol O₂ g^?1 FW h^?1 (μmol m^?2 s^?1)^?1 in the green algal species. At 0°C P_max values of the brown and red algae ranged from 6.8 to 19.1 μmol O₂ g^?1 FW h^?1 and were similarly high or higher than those of comparable Arctic-cold temperate species. Optimum temperatures for photosynthesis were 5 to 10°C in A. mirabilis, 10°C in H. grandifolius, 15°C in G. skottsbergii and 20°C or higher in D. anceps and I. cordata. P: R ratios strongly decreased in most brown and red algae with increasing temperatures due to different Q₁₀ values for photosynthesis (1.4 to 2.5) and dark respiration (2.5 to 4.1). These features indicate considerable physiological adaptation to the prevailing low light conditions and temperatures of Antarctic waters. In this respect the lower depth distribution limits and the northern distribution boundaries of these species partly depend on the physiological properties described here.     

The effects of development at sub-optimal growth temperatures on photosynthetic capacity and susceptibility to chilling-dependent photoinhibition in Zea mays

When plants of Zea mays L. cv. LG11 that have been grown at optimal temperatures are transferred to chilling temperatures (0–12°C) photoinhibition of photosynthetic CO₂ assimilation can occur. This study examines how growth at sub-optimal temperatures alters both photosynthetic capacity and resistance to chilling-dependent photoinhibition. Plants of Z. mays cv. LG11 were grown in controlled environments at 14, 17, 20 and 25°C. As a measure of the capacity for photosynthesis under light limiting conditions, the maximum quantum yields of CO₂ assimilation (φ^a.c) and O₂ evolution (φ^a.o) were determined for the laminae of the second leaves at photon fluxes of 50–150 μmol m^-2s^-1. To determine photosynthetic capacity at photon fluxes approaching light saturation, rates of CO₂ uptake (A₁₅₀₀) and O₂ evolution (A₁₅₀₀) were determined in a photon flux of 1500 μmol m^-2s^-1. In leaves developed at 14°C, φ and φ were 26 and 43%, respectively, of the values for leaves grown at 25°C. Leaves grown at 17°C showed intermediate reductions in φ and φ, whilst leaves developed at 20°C showed no significant differences from those grown at 25°C. Similar patterns of decrease were observed for A₁₅₀₀ and A_1500.0 with decreasing growth temperature. Leaves developed at 25°C showed higher rates of CO₂ assimilation at all light levels and measurement temperatures in comparison to leaves developed at 17 and 14°C. A greater reduction in A₁₅₀₀ relative to A_1500.0 with decreasing growth temperature was attributed to increased stomatal limitation. Exposure of leaves to 800–1000 μmol m^-2 s^-1 when plant temperature was depressed to ca 6.5°C produced a photoinhibition of photosynthetic CO₂ assimilation in all leaves. However, in leaves developed at 17°C the decrease in A₁₅₀₀ following this chilling treatment was only 25% compared to 90% in leaves developed at 25°C. Recovery following chilling was completed earlier in leaves developed at 17°C. The results suggest that growth at sub-optimal temperatures induces increased tolerance to exposure to high light at chilling temperatures. This is offset by the large loss in photosynthetic capacity imposed by leaf development at sub-optimal temperatures.