The responses of photosynthetic gas exchange and chlorophyll a fluorescence to changes of irradiance and temperature in two species of Miscanthus

Photosynthetic CO₂ uptake and chlorophyll (Chl) a fluorescence of C₄ perennial grasses, Miscanthus floridulus (Labill) Warb and M. transmorrisonensis Hayata, from altitudes in central Taiwan of 390 and 2700 m, respectively, were studied at 10 and 25 °C to find if the species differ in their photosynthetic responses to a low temperature, and whether their photosystems 2 become more susceptible to the photoinhibition at low temperatures. For both species, the maximum photosynthetic rate (P_max) was reduced when the leaves were exposed to 10 °C. At irradiances higher than 400 µmol m^-2 s^-1, the values of F_v/F_m were reduced in both species at 10 °C but not at 25 °C, which indicated the photoinhibition at 10 °C. Reductions in P_max and the values of F_v/F_m at 10 °C were lesser in M. transmorrisonensis than in M. floridulus.     

The responses of photosynthetic gas exchange and chlorophyll a fluorescence to changes of irradiance and temperature in two species of Miscanthus

Photosynthetic CO₂ uptake and chlorophyll (Chl) a fluorescence of C₄ perennial grasses, Miscanthus floridulus (Labill) Warb and M. transmorrisonensis Hayata, from altitudes in central Taiwan of 390 and 2700 m, respectively, were studied at 10 and 25 °C to find if the species differ in their photosynthetic responses to a low temperature, and whether their photosystems 2 become more susceptible to the photoinhibition at low temperatures. For both species, the maximum photosynthetic rate (P_max) was reduced when the leaves were exposed to 10 °C. At irradiances higher than 400 μmol m^-2 s^-1, the values of F_v/F_m were reduced in both species at 10 °C but not at 25 °C, which indicated the photoinhibition at 10 °C. Reductions in P_max and the values of F_v/F_m at 10 °C were lesser in M. transmorrisonensis than in M. floridulus. This revised version was published online in June 2006 with corrections to the Cover Date.     

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.     

Prolonging the hydration and active metabolism from light periods into nights substantially enhances lichen growth

This study investigates how hydration during light and dark periods influences growth in two epiphytic old forest lichens, the green algal Lobaria pulmonaria and the cyanobacterial L. scrobiculata. The lichens were cultivated in growth chambers for 14 days (200 μmol m^?1 s^?2; 12 h photoperiod) at four temperature regimes (25/20 °C, 21/16 °C, 13/8 °C, and 6/1 °C; day/night temperatures) and two hydration regimes (12 h day-time hydration; 12 h day-time + 12 h night-time hydration). Growth was highly dynamic, showing that short-term growth experiments in growth cabinets have a high, but largely unexplored potential in functional lichen studies. The highest measured growth rates were not far from the maximal dry matter gain estimated from published net photosynthetic CO₂ uptake data. For the entire data set, photobiont type, temperature, hydration regime and specific thallus mass accounted for 46.6 % of the variation in relative growth rate (RGR). Both species showed substantially higher relative growth rates based on both biomass (RGR) and thallus area (RT_AGR) when they were hydrated day and night compared to hydration in light only. Chronic photoinhibition was substantial in thalli hydrated only during the day time and kept at the highest and lowest temperature regimes, resulting in exponential increases in RGR with increasing maximal PSII efficiency (F _v/F _m) in both species. However, the depression in F _v/F _m was stronger for the cyanolichen than for the cephalolichen at extreme temperatures. The growth-stimulating effect of night-time hydration suggests that nocturnal metabolic activity improves recovery of photoinhibition and/or enhances the conversion rate of photosynthates into thallus extension.     

Influence of nitrogen supply and growth irradiance on photoinhibition and recovery in Heuchera americana (Saxifragaceae)

Prior work demonstrated that Heuchera americana, an evergreen herb inhabiting the deciduous forest understory in the southeastern United States, has a 3-4-fold greater photosynthetic capacity under the low-temperature, strong-light, open canopies of winter compared to the high-temperature, weak-light, closed canopies of summer. Moreover, despite the reductions in soil nitrogen, the chilling temperatures, and the increased quantum flux associated with winter, chronic photoinhibition was not observed in this species at this time of the year. We were interested in the photosynthetic acclimation and photoinhibition characteristics of this species when grown under contrasting light and nitrogen regimes. Newly expanded shade-acclimated leaves of forest-grown plants exposed to strong light varying in intensity and duration at 25°C showed a reduction in F_v/F_m (the ratio of variable to maximum room temperature chlorophyll fluorescence measured after dark adaptation), which was correlated with a decline in ø_a (the intrinsic quantum yield of CO₂-saturated O₂ evolution on an absorbed light basis). Plants grown in the glasshouse under contrasting light (high and low light; HL and LL, respectively) and nitrogen supply (high and low nitrogen; HN and LN, respectively) regimes showed that photosynthetic acclimation to HL was impaired in LN regimes. The HL-LN plants also had the lowest values of F_v/F_m and of ø on both incident and absorbed light bases and had 50% less chlorophyll (per unit area) compared to plants from other growth regimes. Controlled exposure to bright light at low temperatures (2-3°C) for 3 h resulted in a sharp decrease in F_v/F_m (and rise in F_o, the minimum fluorescence yield) in all plants. Shade-grown plants from both N regimes were highly susceptible to chronic photoinhibition, as indicated by a greater reduction in F_v/F_m and incomplete recovery after 18 h in weak light at 25°C. The HL-HN plants were the least susceptible to chronic photoinhibition, having the smallest decrease in F_v/F_m with near full recovery within 6 h. The decline in Fv/Fm in HL-LN plants was comparable to that of shade-acclimated plants, but recovered fully within 6 h. Low-N plants from both light regimes displayed greater increases in F_o which did not return to pretreatment levels after 18 h of recovery. These studies indicate that HL-LN plants were sensitive to chronic photoinhibition and, at the same time, had a high capacity for dynamic photoinhibition. Experimental garden studies showed that H. americana grown in an open field in summer were photoinhibited and did not fully recover overnight or during extended periods of weak light. These results are discussed in relation to the photosynthetic acclimation of H. americana under natural conditions.     

Photosynthetic activity including the thermal‐ and chilling‐light sensitivities of a temperate Japanese brown alga Sargassum macrocarpum

The effects of irradiance, temperature, thermal‐ and chilling‐light sensitivities on the photosynthesis of a temperate alga, Sargassum macrocarpum (Fucales) were determined by a pulse amplitude modulation (PAM)‐chlorophyll fluorometer and dissolved oxygen sensors. Oxygenic photosynthesis–irradiance curves at 8, 20, and 28°C revealed that the maximum net photosynthetic rates (NP _max) and saturation irradiance were highest at 28°C, and lowest at 8°C. Gross photosynthesis and dark respiration determined over a range of temperatures (8–36°C) at 300 μmol photons m^?2 s^?1 revealed that the maximum gross photosynthetic rate (GP_max) occurred at 27.8°C, which is consistent with the highest seawater temperature in the southern distributional limit of this species in Japan. Additionally, the maximum quantum yields of photosystem II (F _v/F _m) during the 72‐h temperature exposures were stable at 8–28°C, but suddenly dropped to zero at higher temperatures, indicative of PSII deactivation. Continuous exposure (12 h) to irradiance of 200 (low) and 1000 (high) μmol photons m^?2 s^?1 at 8, 20, and 28°C revealed greater declines in their effective quantum yields (Φ _PSII) under high irradiance. While Φ _PSII under low irradiance were very similar with the initial F _v/F _m under 20 and 28°C, values rapidly decreased with exposure duration at 8°C. At this temperature, F _v/F _m did not recover to initial values even after 12 h of dark acclimation. Final F _v/F _m of alga at 28°C under high irradiance treatment also did not recover, suggesting its sensitivity to photoinhibition at both low and high temperatures. These photosynthetic characteristics reflect both the adaptation of the species to the general environmental conditions, and its ability to acclimate to seasonal changes in seawater temperature within their geographical range of distribution.     

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.     

The eurythermal adaptivity and temperature tolerance of a newly isolated psychrotolerant Arctic <Emphasis Type="Italic">Chlorella</Emphasis> sp.

A new strain of Chlorella sp. (Chlorella-Arc), isolated from Arctic glacier melt water, was found to have high specific growth rates (μ) between 3 and 27 °C, with a maximum specific growth rate of 0.85 day^?1 at 15 °C, indicating that this strain was a eurythermal strain with a broad temperature tolerance range. To understand its acclimation strategies to low and high temperatures, the physiological and biochemical responses of the Chlorella-Arc to temperature were studied and compared with those of a temperate Chlorella pyrenoidosa strain (Chlorella-Temp). As indicated by declining F _v/F _m, photoinhibition occurred in Chlorella-Arc at low temperature. However, Chlorella-Arc reduced the size of the light-harvesting complex (LHC) to alleviate photoinhibition, as indicated by an increasing Chl a/b ratio with decreasing temperatures. Interestingly, Chlorella-Arc tended to secrete soluble sugar into the culture medium with increasing temperature, while its intracellular soluble sugar content did not vary with temperature changes, indicating that the algal cells might suffer from osmotic stress at high temperature, which could be adjusted by excretion of soluble sugar. Chlorella-Arc accumulated protein and lipids under lower temperatures (<15 °C), and its metabolism switched to synthesis of soluble sugar as temperatures rose. This reflects a flexible ability of Chlorella-Arc to regulate carbon and energy distribution when exposed to wide temperature shifts. More saturated fatty acids (SFA) in Chlorella-Arc than Chlorella-Temp also might serve as the energy source for growth in the cold and contribute to its cold tolerance.     

Two different strategies of Mediterranean macchia plants to avoid photoinhibitory damage by excessive radiation levels during summer drought

The adaptive strategies to high radiation and water stress of the drought tolerant evergreen sclerophylls Quercus coccifera and Arbutus unedo are compared to those of the semi-deciduous Cistus spp. (C. albidus and C. monspeliensis). Cistus spp. partially avoided drought by a marked reduction of their transpirational surface through leaf abscission during summer, when predawn water potential declined below -5.5 MPa. Chlorophyll fluorescence measurements revealed a reversible diurnal decrease of maximum photochemical efficiency of PSII (F_v/F_m), which became more accentuated during summer drought in all species. An important strategy to avoid damage by excessive radiation levels in Cistus spp. was the structural regulation of light interception through leaf angle changes, from a more horizontal orientation in spring (< 35°) to a more vertical orientation in summer (> 70°). Horizontal orientated leaves were highly susceptible to photoinhibition, and excessive radiation often resulted in irreversible photodamage followed by leaf abscission during summer, whereas vertical leaf orientation appeared to protect the leaf from severe photoinhibition. Still, these mechanisms were not fully successful in avoiding chronic photoinhibition, and predawn F_v/F_m values remained low in Cistus spp. during summer (only exhibiting a partial overnight recovery). Evergreen sclerophylls were less susceptible to photoinhibition, and the diurnal decline in F_v/F_m remained fully reversible during drought. Structural regulation of light interception was not found to be an important strategy in these species, and only small, though significant changes in leaf angle occurred. The ecological importance of the adaptive strategies of each functional group is discussed.     

Photosynthetic temperature response of the Antarctic vascular plants Colobanthus quitensis and Deschampsia antarctica

The photosynthetic temperature response of the Antarctic vascular plants Colobanthus quitensis and Deschampsia antarctica was examined by measuring whole-canopy CO₂ gas exchange and chlorophyll (Chl) a fluorescence of plants growing near Palmer Station along the Antarctic Peninsula. Both species had negligible midday net photosynthetic rates (P_n) on warm, usually sunny, days (canopy air temperature [T_c]> 20°C), but had relatively high P_n on cool days (T_c<10°C). Laboratory measurements of light and temperature responses of P_n showed that high temperature, not visible irradiance, was responsible for depressions in P_n on warm sunny days. The optimal leaf temperatures (T_l) for P_n in C. quitensis and D. antarctica were 14 and 10°C, respectively. Both species had substantial positive P_n at 0°C T_l, which were 28 (C. quitensis) and 32% (D. antarctica) of their maximal P_n, and we estimate that their low-temperature compensation points occurred at ?2°C T_l (C. quitensis) and ?3°C (D. antarctica). Because of the strong warming trend along the peninsula over recent decades and predictions that this will continue, we were particularly interested in the mechanisms responsible for their negligible rates of P_n on warm days and their unusually low high-temperature compensation points (i.e., 26°C in C. quitensis and 22°C in D. antarctica). Low P_n at supraoptimal temperature (25°C) appeared to be largely due to high rates of temperature-enhanced respiration. However, there was also evidence for direct impairment of the photosynthetic apparatus at supraoptimal temperature, based on Chl fluorescence and P_n/intercellular CO₂ concentration (c_i) response curve analyses. The breakpoint or critical temperature (T_cr) of minimal fluorescence (F_o) was ≈42°C in both species, which was well above the temperatures where reductions in P_n were evident, indicating that thylakoid membranes were structurally intact at supraoptimal temperatures for P_n. The optimal T_l for photochemical quenching (q_p) and the quantum yield of photosystem II (PSII) electron transfer (φ_PSII) were 9 and 7°C in C. quitensis and D. antarctica, respectively. Supraoptimal temperatures resulted in lower q_p and greater non-photochemical quenching (q_NP), but had little effect on F_o, maximal fluorescence (F_m) or the ratio of variable to maximal fluorescence (F_v/F_m) in both species. In addition, carboxylation efficiencies or initial slopes of their P_n/c_i response were lower at supraoptimal temperatures, suggesting reduced activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco). Although continued warming along the peninsula will increase the frequency of supraoptimal temperatures, T_c at our field site averaged 4.3°C and was below the temperature optima for P_n in these species for the majority of diurnal periods (86%) during the growing season, suggesting that continued warming will usually improve their rates of P_n.     

Photosynthetic response of Nereocystis luetkeana (Phaeophyta) to high light

Photosynthetic response to high light was determined for Bull kelp, Nereocystis luetkeana (K. Mertens) Postels and Ruprecht in order to understand how this species is affected by short‐term fluctuations in irradiance. Exposure of N. luetkeana blades to high intensity photosynthetically active radiation (1000 µmol photons m^?2 s^–1) caused increased non‐photochemical quenching of fluorescence and higher de‐epoxidation ratios for xanthophyll pigments indicating that energy‐quenching xanthophylls were used to protect blades against photoinhibition. Despite initiation of these photoprotective mechanisms, maximum photochemical efficiency of photosystem II (F_v/F_m) decreased 40% in response to a 60 min exposure to 1000 µmol photons m^?2 s^–1 photosynthetically active radiation indicating that photoinhibition had occurred. Light‐saturated rates of oxygen evolution were not changed significantly by the high light treatment. Recovery of maximum photochemical efficiency of photosystem II to within 8% of initial values occurred after a 300‐min dim light period. Younger sections of the blades were slightly more susceptible to high light damage than older sections. Middle sections of the blades were more prone to light‐induced damage at water temperatures of 7°C or 18°C, as compared to 13°C. Exposure to biologically effective ultraviolet‐B radiation (UV‐B_be) (up to 4.5 kJ m^–2 day^–1) in photoinhibitory light conditions did not significantly affect light‐induced damage to photosystem II.     

Temperature-induced changes of malondialdehyde, heat-shock proteins in relation to chlorophyll fluorescence and photosynthesis in Conocarpus lancifolius (Engl.)

The effect of variable temperatures (10–50 °C) on photosynthesis and chlorophyll fluorescence in Conocarpus lancifolius was evaluated. Additionally, the ability of the species to synthesize heat-shock proteins (HSPs) to protect against high temperatures, and malondialdehyde (MDA) as a by-product of lipid peroxidation was investigated. Plants at 10 °C showed virtually no measurable growth, leaf discoloration and a few brown lesions, while high temperatures (40 and 50 °C) promoted growth and lateral branch development. Chlorophyll content index, photochemical efficiency (F _v/F _m) of PS II, electron transport rate and photosynthetic rate declined with decreasing temperature but increased significantly at higher temperatures. Heat-shock protein (HSP 70 kDa) was produced at temperatures 30–50 °C and an additional 90 kDa protein was also produced at 50 °C. Increase in the efficiency of excitation energy captured by the open PS II reaction centers (F _v/F _m) increased linearly (P ≤ 0.05) with the accumulation of HSP 70 at higher temperatures. However, at low temperatures the concentration of MDA increased significantly, indicating lipid peroxidation due to oxidative stress. The production and accumulation of HSP 70 and 90 kDa coupled with increased electron transport rate and photochemical efficiency can be used to assess survival, growth capacity and to some extent the tolerance of C. lancifolius to elevated temperatures.     

Photoinhibition in the Antarctic moss Grimmia antarctici Card when exposed to cycles of freezing and thawing

Freezing and thawing of the endemic moss species Grimmia antarctici Card, caused photoinhibition. When snow cover was removed from moss in the field, resulting in exposure to fluctuating temperatures and light conditions, photoinhibition, measured as a reduction in the ratio of variable to maximum chlorophyll a fluorescence (F_v/F_m), was observed. The extent of photoinhibition was highly variable and appeared to be reversible during periods of warmer temperatures. A series of controlled laboratory studies found that the light conditions that prevail between freezing and thawing events influenced the recovery from photoinhibition observed during freezing and thawing, with low light conditions facilitating the greatest rates of recovery. After four cycles of freezing and thawing, recovery from photoinhibition in hydrated moss was achieved within 12 h of transfer to 5°C and 15 μmol quanta m^?2 s^?1. These results favour the hypothesis that photoinhibition observed during freezing represents a protective process involving the down-regulation of photo-system II when photosynthetic carbon assimilation is limited by low temperatures.     

Effects of photoinhibitory treatment on CO2 assimilation, the quantum yield of CO2 assimilation, D1 protein, ascorbate, glutathione and xanthophyll contents and the electron transport rate in vine leaves

The responses of photosynthesis to high light and low temperature were studied in vines cultivated in the greenhouse in low light. Exposure to high light (1000 /umol m^?2 s^?1) or low temperature (5 °C) alone had no measurable effect on the photosynthetic processes, but the combination of high light and low temperature caused rapid loss of photosynthetic capacity and a decrease in the efficiency of photosynthetic energy conversion. After a 15 h exposure to 5°C at high light, the F_v/sb/F_mratio had decreased by 80% and the apparent quantum yield by 75%. Nevertheless, when the leaves were returned to low light at 22°C, these parameters recovered rapidly. The foliar pools of ascorbate and glutathione decreased in the first hours of photoinhibitory treatment while the zeaxanthin content increased from negligible levels to about 50% of the total foliar xanthophyll pool. There was a clear correlation between the zeaxanthin content of the leaves and their F_v/F_m ratio during both photoinhibition and recovery. However, there was also a good correlation between the decrease in theF_v F_m ratio and the measured decrease in the total foliar levels of the antioxidants ascorbate and glutathione. The amount of D, protein diminished over the same period as the zeaxanthin levels were increasing. This approach, involving simultaneous measurements of several parameters considered to influence photosystemy II activity, clearly demonstrates that measured decreases in F_v/F_m may not simply be related to zeaxanthin levels or to amounts of D₁ protein alone but result from multifactoral influences.     

Carotenoid composition in Zea mays developed at sub-optimal temperature and different light intensities

The content and composition of pigments were examined in the third leaf of Zea mays L. plants grown under controlled environment at near-optimal temperature (24°C) or sub-optimal temperature (14°C) at a light intensity of either 200 or 600 μmol m^?2 s^?1. Compared to leaves grown at 24°C, leaves grown at 14°C showed a large reduction in the chlorophyll (Chl) content, a marked decrease in the Chl a/b ratio, and a large increase in the ratio of total carotenoids/Chl a+b. Leaves grown at 14°C showed a much lower content of β-carotene than leaves grown at 24°C, while the content of the carotenoids of the xanthophyll cycle (violaxanthin [V] + antheraxanthin [A] + zeaxanthin [Z]) was markedly higher in the former leaves as compared to the latter leaves; neoxanthin and lutein were affected by the growth temperature to a much lesser extent. The xanthophylls/β-carotene ratio was about three times higher in leaves grown at 14°C as compared to leaves grown at 24°C. On a chlorophyll basis, the two types of leaves hardly differed in their level of β-carotene, while the levels of the xanthophylls (including lutein and neoxanthin) were higher in 14°C-grown leaves as compared to 24°C-grown leaves. In leaves grown at 14°C, 40 and 56% of the V+A+Z pool was in the form of zeaxanthin at low light intensity and high light intensity, respectively. Only trace amounts of zeaxanthin, if any, were present in leaves grown at 24°C. The changes in the pigment composition induced by growth at sub-optimal temperature were more pronounced at a light intensity of 600 as compared to 200 μmol m^?2 s^?1. In the given range, the light intensity slightly affected the composition of pigments in leaves grown at 24°C. The physiological significance of the modifications to the pigment composition induced by growth at sub-optimal temperature is discussed.     

Cold Acclimation Ability and Photosynthesis among Species of the Tropical Coffea Genus

Abstract: Three Coffea species (C. arabica cv. Icatu, C. canephora cv. Apoatã and C. dewevrei) were tested in order to identify and study the mechanisms of tolerance to low, non‐freezing temperatures. Several photosynthesis‐related parameters were monitored during a 20‐day period of gradual temperature decrease, from 25/20 °C (day/night) down to 15/10 °C, during chilling treatments (15/4 °C), and upon rewarming (25/20 °C). Differences were found among species, both during low temperature exposure and during rewarming. In general, Coffea species showed cold‐induced photoinhibition of photosynthesis, which was attributable to biochemical (in vivo ribulose‐1,5‐bisphosphate carboxylase/oxygenase activity and carbohydrate synthesis) and biophysical (antennae functioning, photosystem II efficiency and linear electron transport) inactivation, rather than to stomatal constraints. The moderately low temperature of 15/10 °C was enough to cause a negative impact on net photosynthesis (A), mostly due to low (initial) rubisco activity in all species. However, C. arabica cv. Icatu showed a higher tolerance to chilling and recovered quickly and completely upon rewarming, as assessed from the impacts on the photosynthetic machinery (e.g. A_max, F_o, F_v/F_m, F_v′/F_m′, q_P, ?_e, rubisco activity) and on carbohydrate metabolism. Such lesser effects are likely to be related to the strong increases and higher contents of zeaxanthin, lutein and β‐carotene that presumably increased the ability to dissipate excitation energy and contributed to protect the photosynthetic apparatus. During cold exposure, a significant reduction of the α/β carotene ratio, which is considered an acclimation feature, was observed solely in C. arabica cv. Icatu. However, C. canephora cv. Apoatã and, especially, C. dewevrei showed to be highly cold‐sensitive. In these latter species, the photoinhibitory impairments to photosynthesis were stronger, probably due to the lower contents of protecting pigments during chilling conditions that lead to a higher vulnerability to excess excitation energy. Moreover, the mesophyll impairments (e.g. A_max, F_v/F_m, ?_e) became significant even at moderately low temperatures of 15/10 °C, and a lower ability to recover after chilling exposure was observed. The limitation of in vivo rubisco activity and A_max may have been due to substrate limitation, but disturbances in sugar metabolism could also play an important role in the expression of chilling sensitivity in C. canephora cv. Apoatã and C. dewevrei.     

The effect of temperature,light-spectrum,desiccation and salinity gradients on the photosynthetic performance of a subtidal brown alga,Sargassum macrocarpum,from Japan

The effect of temperature, light-spectrum, desiccation and salinity gradients on the photosynthesis of a Japanese subtidal brown alga, Sargassum macrocarpum (Fucales), was determined using a pulse amplitude modulation-chlorophyll fluorometer and dissolved oxygen sensors. Temperature responses of the maximum (F_v/F_m in darkness) and effective (ΔF/F_m′ at 50 μmol photons m⁻² s⁻¹; = Φ_PSII) quantum yields during 6-day culture (4–36°C) remained high at 12–28°C, but decreased at higher temperatures. Nevertheless, ΔF/F_m′ also dropped at temperatures below 8°C, suggesting light sensitivity under chilling temperatures because F_v/F_m remained high. Photosynthesis–irradiance responses at 24°C under red (660 nm), green (525 nm), blue (450 nm) and white light (metal halide lamp) showed that maximum net photosynthesis under blue and white light was greater than under red and green light, indicating the sensitivity and photosynthetic availability of blue light in the subtidal light environment. In the desiccation experiment, samples under aerial exposure of up to 8 h under dim-light at 24°C and 50% humidity showed that ΔF/F_m′ quickly declined after more than 45 min of emersion; furthermore, ΔF/F_m′ also failed to recover to initial levels even after 1 day of rehydration in seawater. Under the emersion state, the ΔF/F_m′ remained high when the relative water content (RWC) was greater than 50%; in contrast, it quickly dropped when the RWC was less than 50%. When the RWC was reduced below 50%, ΔF/F_m′ did not return to initial levels, regardless of subsequent re-hydration, suggesting a low capacity of photosynthesis to recover from desiccation. The stenohaline response of photosynthesis under 3-day culture is evident, given that ΔF/F_m′ declined when salinity was beyond 20–40 psu. Adaptation to subtidal environments in temperate waters of Japan can be linked to these traits.     

Response of effective quantum yield of photosystem 2 to <Emphasis Type="Italic">in situ</Emphasis> temperature in three alpine plants

The response of effective quantum yield of photosystem 2 (ΔF/F_m’) to temperature was investigated under field conditions (1 950 m a.s.l.) in three alpine plant species with contrasting leaf temperature climates. The in situ temperature response did not follow an optimum curve but under saturating irradiances [PPFD >800 µìmol(photon) m^?2s^?1] highest ΔF/F_m’ occurred at leaf temperatures below 10°C. This was comparable to the temperature response of antarctic vascular plants. Leaf temperatures between 0 and 15°C were the most frequently (41 to 56%) experienced by the investigated species. At these temperatures, ΔF/F_m’ was highest in all species (data from all irradiation classes included) but the species differed in the temperature at which ΔF/F_m’ dropped below 50% (Soldanella pusilla >20°C, Loiseleuria procumbens >25°C, and Saxifraga paniculata >40°C). The in situ response of ΔF/F_m’ showed significantly higher ΔF/F_m’ values at saturating PPFD for the species growing in full sunlight (S. paniculata and L. procumbens) than for S. pusilla growing under more moderate PPFD. The effect of increasing PPFD on ΔF/F_m’, for a given leaf temperature, was most pronounced in S. pusilla. Despite the broad diurnal leaf temperature amplitude of alpine environments, only in S. paniculata did saturating PPFD occur over a broad range of leaf temperatures (43 K). In the other two species it was half of that (around 20 K). This indicates that the setting of environmental scenarios (leaf temperature×PPFD) in laboratory experiments often likely exceeds the actual environmental demand in the field.     

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.     

Light and temperature dependent inhibition of photosynthesis in frost-hardened and un-hardened seedlings of pine

Needles of un-hardened and frost-hardended seedlings of Pinus sylvestris and Pinus contorta were exposed to photoinhibitory photon flux densities at temperatures between 0 and 35°C under laboratory conditions. Photoinhibition of photosynthesis was assayed by measuring oxygen evolution under saturating CO₂ in a leaf disc oxygen electrode or by recording of photosystem II fluorescence induction kinetics at 77 K. It was demonstrated that frost hardening of pine did not affect the susceptibility of photosynthesis to short time (2 h) photoinhibition at 15°C. The two pine species irrespective of acclimative state were equally sensitive to photoinhibition as assayed by apparent photon yield analyses of photosynthetic oxygen evolution. Plots of the apparent photon yield of oxygen evolution vs. F _v /F _m revealed a non-linear relationship.In the temperature range of 15–20°C short term photoinhibition caused a loss of F _v without effect on F ₀ . However, photoinhibition at temperatures lower or higher caused F ₀ to increase and decrease, respectively. In fact the decrease of F _v v _/F upon lowering the temperature was mainly caused by the temperature effect on F ₀ . Besides photoinhibition causing the well established quenching of F _v by increased radiationless decay somewhere in the reaction center-antenna complex, it is suggested that F ₀ generally increases as a result of loss of functional reaction centers causing decreased trapping of excitation energy. However, the high temperature induced quenching of F ₀ suggests that the quenching process (or processes) induced under photoinhibitory conditions is temperature dependent; i.e. it increases with the increase of temperature.In pine the photon yield of photosynthesis was much more sensitive to short term photoinhibition than was the rate of light saturated photosynthesis. This difference is explained by photosystem II and electron transport having surplus capacity relative to that of reductive carbon metabolism.