INORGANIC CARBON REPLETION CONSTRAINS STEADY‐STATE LIGHT ACCLIMATION IN THE CYANOBACTERIUM SYNECHOCOCCUS ELONGATUS1

Cyanobacteria show high metabolic plasticity by re‐allocating macromolecular resources in response to variations in both environmental inorganic carbon (Ci) and light. We grew cultures of the picoplanktonic cyanobacterium Synechococcus elongatus Nägeli across a 50‐fold range of growth irradiance at either a dissolved [Ci] <0.1 mM, sufficient to induce strongly the carbon‐concentrating mechanism (CCM) or a dissolved [Ci] of ～4 mM, sufficient to strongly induce the CCM to basal constitutive activity. There was no detectable growth cost of acclimation to low Ci across the entire range of irradiance and growth was nearly light saturated at 50 l mol photons·m^?2·s^?1. Cells acclimated to low Ci significantly re‐allocated macromolecular resources to support their CCM, while maintaining near homeostatis of metabolic flux per unit photosynthetic complex. Changing growth irradiance also drove re‐organization of the photosynthetic machinery to balance excitation flux and metabolic demands, but flux per complex varied widely across the range of tolerable growth irradiances. Across the range of growth irradiance, low Ci cells had significantly less phycocyanin than high Ci cells, which corresponded to a lower PSII absorbance capacity. Furthermore, low Ci cells maintained more PSI per cell^?1 than high Ci cells under high growth irradiance. Low Ci cells could therefore maintain more of their PSII reaction centers open at high growth irradiance than could high Ci cells, which experienced a significant PSII closure. Thus, acclimation to growth under high available Ci actually constrained acclimation to high light by restricting electron transport downstream from PSII in S. elongatus.     

CYANOBACTERIAL ACCLIMATION TO RAPIDLY FLUCTUATING LIGHT IS CONSTRAINED BY INORGANIC CARBON STATUS1

Acclimation to rapidly fluctuating light, simulating shallow aquatic habitats, is altered depending on inorganic carbon (C_i) availability. Under steady light of 50 μmol photons·m^?2·s^?1, the growth rate of Synechococcus elongatus PCC7942 was similar in cells grown in high C_i (4 mM) and low C_i (0.02 mM), with induced carbon concentrating mechanisms compensating for low C_i. Growth under fluctuating light of a 1‐s period averaging 50 μmol photons·m^?2·s^?1 caused a drop in growth rate of 28%±6% in high C_i cells and 38%±8% in low C_i cells. In high C_i cells under fluctuating light, the PSI/PSII ratio increased, the PSII absorption cross‐section decreased, and the PSII turnover rate increased in a pattern similar to high‐light acclimation. In low C_i cells under fluctuating light, the PSI/PSII ratio decreased, the PSII absorption cross‐section decreased, and the PSII turnover remained slow. Electron transport rate was similar in high and low C_i cells but in both was lower under fluctuating than under steady light. After acclimation to a 1‐s period fluctuating light, electron transport rate decreased under steady or long‐period fluctuating light. We hypothesize that high C_i cells acclimated to exploit the bright phases of the fluctuating light, whereas low C_i cells enlarged their PSII pool to integrate the fluctuating light and dampen the variation of the electron flux into a rate‐restricted C_i pool. Light response curves measured under steady light, widely used to predict photosynthetic rates, do not properly predict photosynthetic rates achieved under fluctuating light, and exploitation of fluctuating light is altered by C_i status.     

Response of photosynthesis to high light and drought for Arabidopsis thaliana grown under a UV-B enhanced light regime

Arabidopsis thaliana grown in a light regime that included ultraviolet-B (UV-B) radiation (6 kJ m⁻² d⁻¹) had similar light-saturated photosynthetic rates but up to 50% lower stomatal conductance rates, as compared to plants grown without UV-B radiation. Growth responses of Arabidopsis to UV-B radiation included lower leaf area (25%) and biomass (10%) and higher UV-B absorbing compounds (30%) and chlorophyll content (52%). Lower stomatal conductance rates for plants grown with UV-B radiation were, in part, due to lower stomatal density on the adaxial surface. Plants grown with UV-B radiation had more capacity to down regulate photochemical efficiency of photosystem II (PSII) as shown by up to 25% lower φ_PSII and 30% higher non-photochemical quenching of chlorophyll fluorescence under saturating light. These contributed to a smaller reduction in the maximum photochemical efficiency of PSII (F _v/F _m), greater dark-recovery of F _v/F _m, and higher light-saturated carbon assimilation and stomatal conductance and transpiration rates after a four-hour high light treatment for plants grown with UV-B radiation. Plants grown with UV-B were more tolerant to a 12 day drought treatment than plants grown without UV-B as indicated by two times higher photosynthetic rates and 12% higher relative water content. UV-B-grown plants also had three times higher proline content. Higher tolerance to drought stress for Arabidopsis plants grown under UV-B radiation may be attributed to both increased proline content and decreased stomatal conductance. Growth of Arabidopsis in a UV-B-enhanced light regime increased tolerance to high light exposure and drought stress.     

PHYSIOLOGICAL AND PHOTOSYNTHETIC RESPONSES OF SYNECHOCYSTIS AQUATILIS F. AQUATILIS (CYANOPHYCEAE) TO ELEVATED LEVELS OF ZINC1

Physiological and structural changes in cells of Synechocystis aquatilis f. aquatilis acclimated to grow in the presence of high zinc levels (2.20–3.30 mg·L^?1) were investigated. Growth of these cells showed a decreased specific growth rate and final yield of about 60% and 50%, respectively, of the values found for cells grown in the presence of 0.21 mg zinc·L^?1 (control culture). The higher the zinc concentration in the culture medium, the more pronounced the reduction in the chl a content. Regardless of zinc concentration, S. aquatilis possessed three distinct carotenoids. A decrease in carotenoid content accompanied the decrease of chl a, and the proportions of the pigments to each other were not affected by zinc. The photosynthetic performance of cells cultured in the presence of high zinc levels showed a decline in both the apparent photosynthetic efficiency and the photosynthetic maximal rate. In these cells the PSII reaction centers became partially closed, and the electron transport activity around PSII and PSI was reduced to 61% and 38% of the control values, respectively, which may indicate an altered PSII/PSI stoichiometry. In addition, electron micrographs revealed a reduced amount of thylakoid membranes, indicating that acclimation to high zinc levels led to a decrease in the overall number of photosynthetic units. On the other hand, light microscopic observation of negative‐stained cells revealed the presence of a thick mucilaginous layer surrounding the high zinc‐acclimated cells. This extracellular material could retain high amounts of metal ions from the medium, thus providing the Synechocystis cells a mechanism to circumvent toxic levels of zinc.     

Isolate-specific effects of ultraviolet radiation on photosynthesis,growth and mycosporine-like amino acids in the microbial mat-forming cyanobacterium <Emphasis Type="Italic">Microcoleus </Emphasis><Emphasis Type="Italic">chthonoplastes</Emphasis>

Microcoleus chthonoplastes constitutes one of the dominant microorganisms in intertidal microbial mat communities. In the laboratory, the effects of repeated daily exposure to ultraviolet radiation (16:8 light:dark cycle) was investigated in unicyanobacterial cultures isolated from three different localities (Baltic Sea = WW6; North Sea = STO and Brittany = BRE). Photosynthesis and growth were measured in time series (12–15 days) while UV-absorbing mycosporine-like amino acids (MAAs) and cellular integrity were determined after 12 and 3 days exposure to three radiation treatments [PAR (22 μmol photon m⁻² s⁻¹) = P; PAR + UV-A (8 W m⁻²) = PA; PAR + UV-A + UV-B (0.4 W m⁻²) = PAB]. Isolate-specific responses to UVR were observed. The proximate response to radiation stress after 1-day treatment showed that isolate WW6 was the most sensitive to UVR. However, repeated exposure to radiation stress indicated that photosynthetic efficiency (F _v/F _m) of WW6 acclimated to UVR. Conversely, although photosynthesis in STO exhibited lower reduction in F _v/F _m during the first day, the values declined over time. The BRE isolate was the most tolerant to radiation stress with the lowest reduction in F _v/F _m sustained over time. While photosynthetic efficiencies of different isolates were able to acclimate to UVR, growth did not. The discrepancy seems to be due to the higher cell density used for photosynthesis compared to the growth measurement. Apparently, the cell density used for photosynthesis was not high enough to offer self-shading protection because cellular damage was also observed in those filaments under UVR. Most likely, the UVR acclimation of photosynthesis reflects predominantly the performance of the surviving cells within the filaments. Different strategies were observed in MAAs synthesis. Total MAAs content in WW6 was not significantly different between all the radiation treatments. In contrast, the additional fluence of UV-A and UV-B significantly increased MAAs synthesis and accumulation in STO while only UV-B fluence significantly increased MAAs content in BRE. Regardless of the dynamic photosynthetic recovery process and potential UV-protective functions of MAAs, cellular investigation showed that UV-B significantly contributed to an increased cell mortality in single filaments. In their natural mat habitat, M. chthonoplastes benefits from closely associated cyanobacteria which are highly UVR-tolerant due to the production of the extracellular UV-sunscreen scytonemin.     

High-light-like photosynthetic responses of Cucumis sativus leaves acclimated to fluorescent illumination with a high red:far-red ratio: interaction between light quality and quantity

This study evaluated the photosynthetic responses of Cucumis sativus leaves acclimated to illumination from three-band white fluorescent lamps with a high red:far-red (R:FR) ratio (R:FR = 10.5) and the photosynthetic responses of leaves acclimated to metal-halide lamps that provided a spectrum similar to that of natural light (R:FR = 1.2) at acclimation photosynthetic photon flux density (PPFD) of 100 to 700 μmol m^?2 s^?1. The maximum gross photosynthetic rate (P _G) of the fluorescent-acclimated leaves was approximately 1.4 times that of the metal-halide-acclimated leaves at all acclimation PPFDs. The ratio of quantum efficiency of photosystem II (Φ_PSII) of the fluorescent-acclimated leaves to that of the metal-halide-acclimated leaves tended to increase with increasing acclimation PPFD, whereas the corresponding ratios for the leaf mass per unit area tended to decrease with increasing acclimation PPFD. These results suggest that the greater maximum P _G of the fluorescent-acclimated leaves resulted from an interaction between the acclimation light quality and quantity, which was mainly caused by the greater leaf biomass for photosynthesis per area at low acclimation PPFDs and by the higher Φ_PSII as a result of changes in characteristics and distribution of chloroplasts, or a combination of these factors at high acclimation PPFDs.     

The effects of phenotypic plasticity on photosynthetic performance in winter rye, winter wheat and Brassica napus

The contributions of phenotypic plasticity to photosynthetic performance in winter (cv Musketeer, cv Norstar) and spring (cv SR4A, cv Katepwa) rye (Secale cereale) and wheat (Triticum aestivum) cultivars grown at either 20°C [non‐acclimated (NA)] or 5°C [cold acclimated (CA)] were assessed. The 22–40% increase in light‐saturated rates of CO₂ assimilation in CA vs NA winter cereals were accounted for by phenotypic plasticity as indicated by the dwarf phenotype and increased specific leaf weight. However, phenotypic plasticity could not account for (1) the differential temperature sensitivity of CO₂ assimilation and photosynthetic electron transport, (2) the increased efficiency and light‐saturated rates of photosynthetic electron transport or (3) the decreased light sensitivity of excitation pressure and non‐photochemical quenching between NA and NA winter cultivars. Cold acclimation decreased photosynthetic performance of spring relative to winter cultivars. However, the differences in photosynthetic performances between CA winter and spring cultivars were dependent upon the basis on which photosynthetic performance was expressed. Overexpression of BNCBF17 in Brassica napus generally decreased the low temperature sensitivity (Q₁₀) of CO₂ assimilation and photosynthetic electron transport even though the latter had not been exposed to low temperature. Photosynthetic performance in wild type compared to the BNCBF17‐overexpressing transgenic B. napus indicated that CBFs/DREBs regulate not only freezing tolerance but also govern plant architecture, leaf anatomy and photosynthetic performance. The apparent positive and negative effects of cold acclimation on photosynthetic performance are discussed in terms of the apparent costs and benefits of phenotypic plasticity, winter survival and reproductive fitness.     

Acclimation of the diatom Stephanodiscus neoastraea and the cyanobacterium Planktothrix agardhii to simulated natural light fluctuations

Functional and structural characteristics of the photosynthetic apparatus were studied in the diatom Stephanodiscus neoastraea and the cyanobacterium Planktothrix agardhii which were grown semi-continuously under constant irradiance or under simulated natural light fluctuations. The light fluctuations consisted of 24 oscillations of exponentially increasing and decreasing irradiance over a 12-h light period. Maximum irradiance was 1100 μmol photons m⁻² s⁻¹ with the ratio of maximum to minimum intensities being 100, simulating Langmuir circulations with a ratio of euphotic to mixing depth of 1. S. neoastraea acclimated to the light fluctuations by doubling the number and halving the size of photosynthetic units (PS II) while the amount of chlorophylls and carotenoids remained unchanged. The chlorophyll-specific maximum photosynthetic rate was enhanced while the slope of the photosynthesis versus irradiance curves was not influenced by the light fluctuations. Acclimation of P. agardhii was mainly characterized by an increase in chlorophyll content. Both photosystems showed only little changes in number and size. Maximum photosynthetic rate, saturating irradiance and initial slope of the photosynthesis versus irradiance curves did not vary. Although both high and low light were contained in the fluctuating light, an analogy to low or high light acclimation was not found for the diatom nor for the cyanobacterium acclimated to light fluctuations. We suggest that the acclimation to fluctuating light is a response type outside the known scheme of low and high light acclimation. This revised version was published online in June 2006 with corrections to the Cover Date.     

The interaction of visible and UV-B light during photodamage and repair of Photosystem II

In order to understand the mechanism of photodamage induced by solar radiation under natural conditions, we studied the interaction of visible and ultraviolet-B light in the inactivation and repair of the Photosystem II complex by using oxygen evolution and flash-induced chlorophyll fluorescence measurements. In isolated spinach thylakoids and Synechocystis 6803 cells, in which de novo protein synthesis is blocked by lincomycin, photodamage of Photosystem II by visible and UV-B light is characterized by linear semilogarithmic inactivation curves for both separate and combined illumination protocols. The extent of PS II inactivation obtained after combined illumination can be well simulated by assuming independent damaging events induced by visible and UV-B photons. In intact Synechocystis cells capable of protein repair, simultaneous illumination by visible and UV-B light impairs Photosystem II activity to a smaller extent than expected from the independent damaging events. This protective effect is pronounced at low visible light (130 μE m⁻² s⁻¹), but becomes negligible at high intensities (1300 μE m⁻² s⁻¹). Exposure of intact Synechocystis 6803 cells to direct sunlight leads to a rapid inactivation of PS II, accompanied by the accumulation of donor side inhibited centers. This phenomenon, which shows the impairment of the manganese cluster of water oxidation was not observed when the ultraviolet components of sunlight were filtered out. We conclude that visible and UV-B photons inactivate PS II via non-interacting mechanisms, which affect different target sites. In intact cells, the two spectral regions do interact, and results in synergistically enhanced protein repair capacity when UV-B radiation is accompanied by low intensity visible light, which provides protection against photodamage. However, this ameliorating effect becomes insignificant at high light intensities characteristic of direct sunlight. This revised version was published online in June 2006 with corrections to the Cover Date.     

Studies on the effects of UV-B radiation on phytoplankton of sub-antarctic lakes and ponds

Summary Experiments were performed to determine the effects of UV-B (ultraviolet, 280–320 nm) radiation on motility and growth of phytoplankton from lakes and ponds in South Georgia. After 4 h of solar radiation and 4h artificial radiation (UV-B_BE 11.6 kJ m^-2 day^-1, UV-B lamps) the swimming velocity of Cryptomonas sp. decreased. The growth rate of Botryococcus, Lyngbya sp. and Stauraslrum sp. did not show any significant variations between the different light conditions. The UV-B component was reduced by filtering solar radiation through glass bottles und cellulose acetate. Cloudy days had only 30% of the radiation of clear days in both the PAR (photosynthetic active radiation) and UV-B regions. The ponds contained large amounts of humic substances, which are responsible for the absorbance in the UV region.     

The ATP-dependent Clp protease is essential for acclimation to UV-B and low temperature in the cyanobacterium Synechococcus

ClpP is the proteolytic subunit of the ATP-dependent Clp protease in eubacteria, mammals and plant chloroplasts. Cyanobacterial ClpP protein is encoded by a multigene family, producing up to four distinct isozymes. We have examined the importance of the first ClpP protein (ClpP1) isolated from the cyanobacterium Synechococcus sp. PCC 7942 for acclimation to ecologically relevant UV-B and low-temperature regimens. When the growth light of 50 μmol photons m^?2 s^?1 was supplemented with 0.5 W m^?2 UV-B for 8 h, the constitutive level of ClpP1 rose eightfold after an initial lag of 1 h. Wild-type cells readily acclimated to this UV-B level, recovering after the initial stress to almost the same growth rate as that before UV-B exposure. Growth of a clpP1 null mutant (ΔclpP1), however, was severely inhibited by UV-B, being eight times slower than the wild type after 8 h. In comparison, ClpP1 content increased 15-fold in wild-type cultures shifted from 37°C to 25°C for 24 h. Wild-type cultures readily acclimated to 25°C after 24 h, whereas the ΔclpP1 strain did not and eventually lost viability with prolonged cold treatment. During acclimation to either UV-B or cold, photosynthesis in the wild type was initially inhibited upon the shift but then recovered. Photosynthesis in ΔclpP1 cultures, however, was more severely inhibited by the stress treatment and failed to recover. Acclimation was also monitored by examining the exchange of photosystem II reaction centre D1 proteins that occurs in wild-type Synechococcus during conditions of excitation stress. During both cold and UV-B shifts, wild-type cultures replaced the acclimative form of D1 (D1:1) with the alternative D1 form 2 (D1:2) within the first hours. Once acclimated to either 25°C or 0.5 W m^?2 UV-B, D1:2 was exchanged back for D1:1. In ΔclpP1 cultures, this second exchange between D1 forms did not occur, with D1:2 remaining the predominant D1 form. Our results demonstrate that the ATP-dependent Clp protease is an essential component of the cold and UV-B acclimation processes of Synechococcus.     

Non-photochemical quenching-dependent acclimation and thylakoid organization of Chlamydomonas reinhardtii to high light stress

Light is essential for all photosynthetic organisms while an excess of it can lead to damage mainly the photosystems of the thylakoid membrane. In this study, we have grown Chlamydomonas reinhardtii cells in different intensities of high light to understand the photosynthetic process with reference to thylakoid membrane organization during its acclimation process. We observed, the cells acclimatized to long-term response to high light intensities of 500 and 1000 µmol m^?2 s^?1 with faster growth and more biomass production when compared to cells at 50 µmol m^?2 s^?1 light intensity. The ratio of Chl a/b was marginally decreased from the mid-log phase of growth at the high light intensity. Increased level of zeaxanthin and LHCSR3 expression was also found which is known to play a key role in non-photochemical quenching (NPQ) mechanism for photoprotection. Changes in photosynthetic parameters were observed such as increased levels of NPQ, marginal change in electron transport rate, and many other changes which demonstrate that cells were acclimatized to high light which is an adaptive mechanism. Surprisingly, PSII core protein contents have marginally reduced when compared to peripherally arranged LHCII in high light-grown cells. Further, we also observed alterations in stromal subunits of PSI and low levels of PsaG, probably due to disruption of PSI assembly and also its association with LHCI. During the process of acclimation, changes in thylakoid organization occurred in high light intensities with reduction of PSII supercomplex formation. This change may be attributed to alteration of protein–pigment complexes which are in agreement with circular dichoism spectra of high light-acclimatized cells, where decrease in the magnitude of psi-type bands indicates changes in ordered arrays of PSII–LHCII supercomplexes. These results specify that acclimation to high light stress through NPQ mechanism by expression of LHCSR3 and also observed changes in thylakoid protein profile/supercomplex formation lead to low photochemical yield and more biomass production in high light condition.

     

Protective mechanisms and acclimation to solar ultraviolet-B radiation in Oenothera stricta

Abstract Mechanisms of plant protection and acclimation to potentially damaging solar ultraviolet-B (UV-B, 280–320 nm) radiation incident on the Earth's surface were examined in Oenothera stricta. Attenuation of this radiation in the upper leaf epidermis reduces the penetration of UV-B radiation to the mesophyll where damage to physiologically sensitive targets can occur. The epidermis is a highly selective radiation filter that can attenuate up to 95% of the incident UV-B radiation and yet transmit between 70% and 80% of the visible radiation. Exposure to UV-B radiation significantly reduced the degree of epidermal UV-B transmittance by as much as 33%. No significant reduction in epidermal transmittance of visible radiation was observed as a result of UV-B exposure. The plasticity in epidermal UV-B transmittance results from production of flavonoid and related phenolic compounds in the tissue. Absorbance of UV-B radiation in llavonoid extract solutions from epidermal and mesophyll tissues significantly increased by as much as 100% and 35%, respectively, after exposure to UV-B radiation. Photosynthetic rates of leaves exposed to UV-B radiation were not significantly reduced at dose rates representative of the radiation flux found in the habitat of this species, but significant photosynthetic depression was observed at dose rates that exceed the field UV-B flux. The phenotypic plasticity in epidermal UV-B transmittance resulting in decreased penetration of damaging UV-B radiation to the mesophyll may reduce the rate of damage to a level where repair mechanisms can keep pace with reduced injury.     

Effects of increasing UV-B radiation and atmospheric CO2 on photosynthesis and growth: implications for terrestrial ecosystems

Increases in UV-B radiation reaching the earth as a result of stratospheric ozone depletion will most likely accompany increases in atmospheric CO₂ concentrations. Many studies have examined the effects of each factor independently, but few have evaluated the combined effects of both UV-B radiation and elevated CO₂. In general the results of such studies have shown independent effects on growth or seed yield. Although interspecific variation is large, high levels of UV-B radiation tends to reduce plant growth in sensitive species, while CO₂ enrichment tends to promote growth in most C₃ species. However, most previous studies have not looked at temporal effects or at the relationship between photosynthetic acclimation to CO₂ and possible photosynthetic limitations imposed by UV-B radiation. Elevated CO₂ may provide some protection against UV-B for some species. In contrast, UV-B radiation may limit the ability to exploit elevated CO₂ in other species. Interactions between the effects of CO₂ enrichment and UV-B radiation exposure have also been shown for biomass allocation. Effects on both biomass allocation and photosynthetic acclimation may be important to ecosystem structure in terms of seedling establishment, competition and reproductive output. Few studies have evaluated ecosystem processes such as decomposition or nutrient cycling. Interactive effects may be subtle and species specific but should not be ignored in the assessment of the potential impacts of increases in CO₂ and UV-B radiation on plants.     

INFLUENCE OF ULTRAVIOLET RADIATION ON GROWTH AND PHOTOSYNTHESIS OF TWO COLD OCEAN DIATOMS1

The influence of chronic exposure to UV-B and UV-A radiation on growth and photosynthesis of two polar marine diatoms (Pseudonitzschia seriata and Nitzschia sp.) was investigated in cultures exposed to moderate photon fluences for 3–7 days. Population growth rates were diminished 50% by UV-B. Fluorescence induction kinetics of photo-system II (PSII) revealed that UV-B caused lower F_v/F_m ratios and half-rise times, indicating damage to the reaction center of PSII and to related elements of the photosynthetic electron transport chain. Carbon assimilation rates per cell and per chlorophyll a were nonetheless highest for UV-B—exposed populations, which also had the highest chlorophyll a content per cell. The UV-B—exposed cells were, however, more vulnerable to visible light-induced photoinhibition. Exposure to UV-A in the absence of UV-B had little effect on growth, fluorescence induction of PSII, or chlorophyll a contents but did have some inhibitory effects on carbon assimilation per chlorophyll a and per cell. The increased photosynthetic capacity of UV-B-exposed cells suggested some ability to compensate for damage to the photosynthetic apparatus.     

The long-term response to fluctuating light quality is an important and distinct light acclimation mechanism that supports survival of <Emphasis Type="Italic">Arabidopsis thaliana</Emphasis> under low light conditions

The long-term response (LTR) of higher plants to varying light qualities increases the photosynthetic yield; however, the benefit of this improvement for physiology and survival of plants is largely unknown, and its functional relation to other light acclimation responses has never been investigated. To unravel positive effects of the LTR we acclimated Arabidopsis thaliana for several days to light sources, which preferentially excite photosystem I (PSI) or photosystem II (PSII). After acclimation, plants revealed characteristic differences in chlorophyll fluorescence, thylakoid membrane stacking, phosphorylation state of PSII subunits and photosynthetic yield of PSII and PSI. These LTR-induced changes in the structure, function and efficiency of the photosynthetic machinery are true effects by light quality acclimation, which could not be induced by light intensity variations in the low light range. In addition, high light stress experiments indicated that the LTR is not involved in photoinhibition; however, it lowers non-photochemical quenching (NPQ) by directing more absorbed light energy into photochemical work. NPQ in turn is not essential for the LTR, since npq mutants performed a normal acclimation. We quantified the beneficial potential of the LTR by comparing wild-type plants with the LTR-deficient mutant stn7. The mutant exhibited a decreased effective quantum yield and produced only half of seeds when grown under fluctuating light quality conditions. Thus, the LTR represents a distinct acclimation response in addition to other already known responses that clearly improves plant physiology under low light conditions resulting in a pronounced positive effect on plant fitness.     

Inhibition of photosynthesis by UV-B exposure and its repair in the bloom-forming cyanobacterium <Emphasis Type="Italic">Microcystis aeruginosa</Emphasis>

The photosynthetic performance of Microcystis aeruginosa FACHB 854 during the process of UV-B exposure and its subsequent recovery under photosynthetic active radiation (PAR) was investigated in the present study. Eight hours UV-B radiation (3.15 W m⁻²) stimulated the increase of photosynthetic pigments content at the early stage of UV-B exposure followed by a significant decline. It suggested that UV-B damage was not an immediate process, and there existed a dynamic balance between damage and adaptation in the exposed cells. Short-term UV-B exposure severely inhibited the photosynthetic capability, but it could restore quickly after being transferred to PAR. Further investigations revealed that the PS II of M. aeruginosa FACHB 854 was more sensitive to UV-B exposure than PS I, and the oxygen-evolving complex of PS II was an important damage target of UV-B. The inhibition of photosynthetic performance caused by UV-B could be recovered to 90.9% of pretreated samples after 20 h exposure at low PAR, but it could not be recovered in the dark as well as under low PAR in the presence of Chloromycetin. It can be concluded that PAR and de novo protein synthesis were essential for the recovery of UV-B-damaged photosynthetic apparatus.     

Effects of physiological integration on photosynthetic efficiency of <Emphasis Type="Italic">Trifolium repens</Emphasis> in response to heterogeneous UV-B radiation

Several studies have found the photosynthetic integration in clonal plants to response to resource heterogeneity, while little is known how it responses to heterogeneity of UV-B radiation. In this study, the effects of heterogeneous UV-B radiation (280–315 nm) on gas exchange and chlorophyll fluorescence of a clonal plant Trifolium repens were evaluated. Pairs of connected and severed ramets of the stoloniferous herb T. repens were grown under the homogeneity (both of ramets received only natural background radiation, ca. 0.6 kJ m⁻² d⁻¹) and heterogeneity of UV-B radiation (one of the ramet received only natural background radiation and the other was exposed to supplemental UV-B radiation, 2.54 kJ m⁻² d⁻¹) for seven days. Stomatal conductance (g _s), intercellular CO₂ concentration (C _i) and transpiration rate (E) showed no significant differences in connected and severed ramets under homogenous and heterogeneous UV-B radiation, however, net photosynthetic rate (P _N) and maximum photosynthetic rate (P _max) of ramets suffered from supplemental increased UV-B radiation and that of its connected sister ramet decreased significantly. Moreover, additive UV-B radiation resulted in a notable decrease of the minimal fluorescence of dark-adapted state (F_o), the electron transport rate (ETR) and photochemical quenching coefficient (q_P) and an increase of nonphotochemical quenching (NPQ) under supplemental UV-B radiation, while physiological connection reverse the results. In all, UV-B stressed ramets could benefit from unstressed ramets by physiological integration in photosynthetic efficiency, and clonal plants are able to optimize the efficiency to maintain their presence in less favourable sites.     

Combined effects of CO2 concentration and enhanced UV-B radiation on faba bean

Due to anthropogenic influences, both solar UV-B irradiance at the earth's surface and atmospheric [CO₂] are increasing. To determine whether effects of CO₂ enrichment on faba bean (cv. Minica) growth are modified by UV-B radiation, the effects of enhanced [CO₂] on growth and photosynthetic characteristics, were studied at four UV-B levels. Faba bean was sensitive to enhanced UV-B radiation as indicated by decreases in total biomass production. Growth stimulation by CO₂ enrichment was greatly reduced at the highest UV-B level. [CO₂] by UV-B interactions on biomass accumulation were related to loss of apical dominance. Both [CO₂] and UV-B radiation affected biomass partitioning, UV-B effects being most pronounced. Effects of [CO₂] and UV-B on faba bean growth were time-dependent, indicating differential sensitivity of developmental stages. [CO₂] and UV-B effects on photosynthetic characteristics were rather small and restricted to the third week of treatment. CO₂ enrichment induced photosynthetic acclimation, while UV-B radiation decreased light-saturated photosynthetic rate. It is concluded that the reduction in biomass production cannot be explained by UV-B-induced effects on photosynthesis.