Chilling-Induced Photoinhibition in Two Oak Species: Are Evergreen Leaves Inherently Better Protected Than Deciduous Leaves?

We compared the sensitivity to cold stress, in terms of photosynthetic capacity and changes in chlorophyll fluorescence of photosystem 2 (PS2), of an evergreen and a deciduous oak species, which co-occur in the southeastern United States. We predicted that the evergreen species, Quercus virginiana, which must endure winter, is likely to have an inherently greater capacity for energy dissipation and to be less susceptible to chilling stress than the deciduous species, Quercus michauxii. Short-term cold stress in both species lead to greater than 50 % reduction in maximum photosynthetic rates, 60-70 % reduction in electron transport, and irreversible quenching of PS2 fluorescence. The kinetics of recovery in the dark after exposure to 1 h high irradiance (1000 μmol m^-2 s^-1) and chilling (5 °C) showed that the evergreen Q. virginiana exhibited more protective q_E and less irreversible quenching (q_I) than the deciduous Q. michauxii. The large q_E which we observed in Q. virginiana suggests that the capacity for photoprotection at low temperatures is not induced by a long-term acclimation to cold but preexists in evergreen leaves. This capacity may contribute to the ability of this species to maintain leaves during the winter. This revised version was published online in June 2006 with corrections to the Cover Date.     

Enhanced photosystem 2 thermostability during leaf growth of Elm (Ulmus pumila) seedlings

We examined photosynthetic activities and thermostability of photosystem 2 (PS2) in leaves of elm (Ulmus pumila) seedlings from initiation to full expansion. During leaf development, net photosynthetic rate (P _N) increased gradually and reached the maximum when leaves were fully developed. In parallel with the increase of P _N, chlorophyll (Chl) content was significantly elevated. Chl a fluorescence measurements showed that the maximum quantum yield of PS2 (ϕ_PS2), the efficiency a trapped exciton, moved an electron into the electron transport chain further than Q_A ⁻ (Ψ_o), and the quantum yield of electron transport beyond Q_A (ϕ_Eo) increased gradually. These results were independently confirmed by our low irradiance experiments. When subjected to progressive heat stress, the young leaves exhibited considerably lower ϕ_PS2 and higher minimal fluorescence (F₀) than the mature leaves, revealing the highly sensitive nature of PS2 under heat in the newly initiating leaves. Further analysis showed that PS2 structure in the newly initiating leaves was strongly altered under heat, as evidenced by the increased fluorescence signals at the position of the K step. We therefore demonstrated an inhibition in the oxygen-evolving complex (OEC) in the young leaves. This resulted in decrease in amount of the functional PS2 reaction centres and relative increase in the PS2 reaction centres with inhibited electron transport at the acceptor side under heat. We suggest that the enhanced thermostability of PS2 during leaf development is associated with improved OEC stability.     

Time-course of photosynthetic induction in four tropical woody species grown in contrasting irradiance habitats

We investigated the photosynthetic induction time-course in species of different ecological groups grown in contrasting forest irradiance environments, gap and understorey, exposed to different darkness times in order to verify the plant capacity to exploit irradiance heterogeneity. Photosynthetic induction was studied in leaves of Bauhinia forficata and Guazuma ulmifolia (early succession species, ES), and Esenbeckia leiocarpa and Hymenaea courbaril (late succession species, LS). T₅₀ and T₉₀ (time estimates to attain 50 and 90 % of maximum net photosynthetic rate, respectively) varied according to the time of previous exposure to darkness and growth irradiance. In both darkness times of 10 and 30 min, T₅₀ was lower in the LS-than ES-species. These results, jointly with significant higher induction state of the leaves after 10 min of darkness, suggest that the LS-species has a higher potential to sunfleck utilization compared to ES-species, both grown in the understorey. After 10 and 30 min of darkness the differences between ecological groups were not clearly detected in the gap for T₅₀ and T₉₀, indicating that eco-physiological characteristics of each ecological group did not influence the induction time of the species evaluated herein. Thus the capacity to show phenotypic plasticity is not exclusive to an ecological group, but it is rather a more intrinsic feature related to the differential capacity of individuals.     

Coal Dust Pollution Effects on Wetland Tree Species in Richards Bay,South Africa

In this study, the effects of coal dust on four, sympatric, wetland tree species in Richards Bay Harbour were investigated. We tested the hypothesis that leaf micromorphology influenced dust accumulation and that coal dust occluded stomata and reduced photosynthetic performance of three mangroves, Avicennia marina, Bruguiera gymnorrhiza and Rhizophora mucronata, and a mangrove associate, Hibiscus tiliaceus. To investigate leaf micromorphology, leaf blade material of the four species was prepared following standard procedures and viewed under scanning electron microscopy. Gas exchange and chlorophyll fluorescence measurements were made at saturating light (>1000 μmol m⁻² s⁻¹) and high temperature (>25 °C) on leaves that were either covered or uncovered with coal dust. There was no evidence of occlusion of stomata by dust. Dust accumulation in A. marina and H. tiliaceus was exacerbated by the presence of a dense mat of trichomes on the undersurface of the leaves, as well as by the sticky brine secreted by salt glands in the former species.Coal dust significantly reduced CO₂ exchange, Photosystem II (PS II) quantum yield and electron transport rate (ETR) through PS II in A. marina and H. tiliaceus but not in the other two mangroves. Reduction in photosynthetic performance was attributed to reduction in light energy incident on the photosynthetic tissues.     

Species-specific acclimation to strong shade modifies susceptibility of conifers to photoinhibition

Photoinhibitory processes in the photosynthetic apparatus of the seedlings of Abies alba (Mill.), Picea abies (Karst.), and Pinus mugo (Turra) growing under strong shade (5 % of full solar irradiance) or full irradiance conditions were investigated in winter and spring using chlorophyll a fluorescence techniques. The extent of photoinhibition in needles as indicated by a decrease in maximum quantum yield of PS II photochemistry (F_v/F_m) depended on species, air temperature and acclimation to the light environment. Unexpectedly, shade-tolerant Abies alba was less affected by low-temperature photoinhibition compared to the other species. F_v/F_m recovered with increasing air temperature. During winter, the seedlings of Picea abies growing in shade showed higher F_v/F_m than those from full light. Non-photochemical quenching of fluorescence (NPQ) measured at the same levels of actinic light was higher in needles acclimated to full light except for Abies alba in February. Photosynthetic performance in term of ETR (apparent electron transfer rate) was also higher in full light-acclimated needles. In April, at ambient temperature, recovery of PS II efficiency from the stress induced by illumination with saturating light was faster in the needles of Picea abies than in those of Abies alba. The shade-acclimated needles of Abies alba and Picea abies showed greater down-regulation of PS II induced by high light stress.     

Photosynthetic rate,chlorophyll fluorescence parameters,and lipid peroxidation of maize leaves as affected by zinc deficiency

Pot trial in greenhouse was conducted using cumulic cinnamon soil from North China to study the effects of zinc deficiency on CO₂ exchange, chlorophyll fluorescence, the intensity of lipid peroxidation, and the activity of superoxide dismutase (SOD) in leaves of maize seedlings. Zn deficiency resulted in a reduction of net photosynthetic rate and stomatal conductance to H₂O. The maximum quantum efficiency of photosystem 2 (PS2) and the PS2 activity were depressed, while the pool size of the plastoquinone molecules was not affected by Zn deficiency. The content of super oxygen anion radical (O₂ ^·−) and the intensity of lipid peroxidation as assessed by malonyldialdehyde content in Zn-deficient leaves were higher than those in Zn-sufficient leaves. The activity of SOD increased with Zn application. The adverse influence of Zn-deficiency on the light stage of photosynthesis is probably one of possible reasons for the limitation of photosynthetic capacity in maize leaves.     

Diurnal changes in chlorophyll fluorescence and light utilization in Colocasia esculenta leaves grown in marshy waterlogged area

Diurnal cycle of chlorophyll fluorescence parameters was done in Colocasia esculenta L. (swamp taro) grown in marshy land under sun or under shade. The sun leaves maintained higher electron transport rate (ETR) and steady state to initial fluorescence ratio (F_s/F₀) than shade leaves. In spite of lower ETR, higher photochemical quenching (PQ), and effective quantum yield of photosystem 2 (Φ_PS2) was evident in shade plants compared to plants exposed to higher irradiance. ETR increased linearly with increase in irradiance more under low irradiance (r ² = 0.84) compared to higher irradiance (r ² = 0.62). The maximum quantum yield of PS 2 (F_v/F_m) did not differ much in sun and shade leaves with the exception of midday when excess of light energy absorbed by plants under sun was thermally dissipated. Hence swamp taro plants adopted different strategies to utilize radiation under different irradiances. At higher irradiance, there was faster decline in proportion of open PS 2 centers (PQ) and excess light energy was dissipated through non-photochemical quenching (NPQ). Under shade, absorbed energy was effectively utilized resulting in higher Φ_PS2.     

Co<Subscript>2</Subscript> assimilation and chlorophyll fluorescence in green <Emphasis Type="Italic">versus</Emphasis> red <Emphasis Type="Italic">Berberis thunbergii</Emphasis> leaves measured with different quality irradiation

Photosynthesis, photorespiration, and chlorophyll (Chl) fluorescence in green and red Berberis thunbergii leaves were studied with two different measuring radiations, red (RR) and “white” (WR). The photosynthetic and photorespiration rates responded differently to the different radiation qualities, which indicate that the carboxylase and oxygenase activities of ribulose-1,5-bisphosphate carboxylase/oxygenase (RuBPCO) were affected. Differences in photosynthetic rate between the two color leaves were less under RR than under WR. However, this reduced difference in photosynthetic rate was not correlated with the stomatal response to the measuring radiation qualities. Compared with the WR, the RR reduced the differences in dark-adapted minimum and maximum fluorescence, steady-state fluorescence, light-adapted maximum fluorescence, and actual photochemical efficiency (Φ_PS2) of photosystem 2 (PS2), but enlarged the difference in non-photochemical quenching between the two color leaves. Differences in both maximum quantum yield of PS2 and ratio of Φ_PS2 to quantum yield of CO₂ fixation between the two color leaves were similar under the two measuring radiations. To exclude disturbance of radiation attenuation caused by anthocyanins, it is better to use RR to compare the photosynthesis and Chl fluorescence in green versus red leaves.     

Light acclimation maintains the redox state of the PS II electron acceptor Q2 within a narrow range over a broad range of light intensities

Chrysanthemum inducum-hybrid `Coral Charm', Hibiscus rosa-sinensis L. `Cairo Red' and Spathiphyllum wallisii Regel `Petit' were grown in natural light in a greenhouse at three levels of irradiance using permanent shade screens. Light acclimation of photosynthesis was characterized using modulated chlorophyll a fluorescence of intact leaves. A close correlation was found between the degree of reduction of the primary electron acceptor Q_A of Photosystem II (PS II) approximated as the fluorescence parameter 1−q_P, and light acclimation. The action range of 1−q_P was 0–0.4 from darkness to full irradiance around noon, within the respective light treatments in the greenhouse, indicating that most PS II reaction centres were kept open. In general, the index for electron transport (ETR) measured by chlorophyll fluorescence was higher for high-light (HL) than intermediate-(IL) and low-light (LL) grown plants. However, HL Chrysanthemum showed 40% higher ETR than HL Hibiscus at light saturation, despite identical redox states of Q_A. The light acclimation of the non-radiative dissipation of excess energy in the antenna, NPQ, varied considerably between the species. However, when normalized against q_P, a strong negative correlation was found between thermal dissipation and ETR measured by chlorophyll fluorescence. To be able to accommodate a high flux of electrons through PS II, the plants with the highest light-saturated ETR had the lowest NPQ/q_P. The possibility of using chlorophyll fluorescence for quantification of the energy balance between energy input and utilization in PS II in intact leaves is discussed. This revised version was published online in June 2006 with corrections to the Cover Date.     

Ambient UV-B radiation reduces PSII performance and net photosynthesis in high Arctic Salix arctica

Ambient ultraviolet-B (UV-B) radiation potentially impacts the photosynthetic performance of high Arctic plants. We conducted an UV-B exclusion experiment in a dwarf shrub heath in NE Greenland (74°N), with open control, filter control, UV-B filtering and UV-AB filtering, all in combination with leaf angle control. Two sites with natural leaf positions had ground angles of 0° (‘level site’) and 45° (‘sloping site’), while at a third site the leaves were fixed in an angle of 45° to homogenize the irradiance dose (‘fixed leaf angle site’). The photosynthetic performance of the leaves was characterized by simultaneous gas exchange and chlorophyll fluorescence measurements and the PSII performance through the growing season was investigated with fluorescence measurements. Leaf harvest towards the end of the growing season was done to determine the specific leaf area and the content of carbon, nitrogen and UV-B absorbing compounds. Compared to a 60% reduced UV-B irradiance, the ambient solar UV-B reduced net photosynthesis in Salix arctica leaves fixed in the 45° position which exposed leaves to maximum natural irradiance. Also a reduced Calvin Cycle capacity was found, i.e. the maximum rate of electron transport (J_max) and the maximum carboxylation rate of Rubisco (V_cmax), and the PSII performance showed a decreased quantum yield and increased energy dissipation. A parallel response pattern and reduced PSII performance at all three sites indicate that these responses take place in all leaves across position in the vegetation. These findings add to the evidence that the ambient solar UV-B currently is a significant stress factor for plants in high Arctic Greenland.     

How to correctly determine the different chlorophyll fluorescence parameters and the chlorophyll fluorescence decrease ratio R<Subscript>Fd</Subscript> of leaves with the PAM fluorometer

This contribution is a practical guide to the measurement of the different chlorophyll (Chl) fluorescence parameters and gives examples of their development under high-irradiance stress. From the Chl fluorescence induction kinetics upon irradiation of dark-adapted leaves, measured with the PAM fluorometer, various Chl fluorescence parameters, ratios, and quenching coefficients can be determined, which provide information on the functionality of the photosystem 2 (PS2) and the photosynthetic apparatus. These are the parameters F_v, F_m, F₀, F_m′, F_v′, NF, and ΔF, the Chl fluorescence ratios F_v/F_m, F_v/F₀, ΔF/F_m′, as well as the photochemical (q_P) and non-photochemical quenching coefficients (q_N, q_CN, and NPQ). q_N consists of three components (q_N = q_E + q_T + q_I), the contribution of which can be determined via Chl fluorescence relaxation kinetics measured in the dark period after the induction kinetics. The above Chl fluorescence parameters and ratios, many of which are measured in the dark-adapted state of leaves, primarily provide information on the functionality of PS2. In fully developed green and dark-green leaves these Chl fluorescence parameters, measured at the upper adaxial leaf side, only reflect the Chl fluorescence of a small portion of the leaf chloroplasts of the green palisade parenchyma cells at the upper outer leaf half. Thus, PAM fluorometer measurements have to be performed at both leaf sides to obtain information on all chloroplasts of the whole leaf. Combined high irradiance (HI) and heat stress, applied at the upper leaf side, strongly reduced the quantum yield of the photochemical energy conversion at the upper leaf half to nearly zero, whereas the Chl fluorescence signals measured at the lower leaf side were not or only little affected. During this HL-stress treatment, q_N, q_CN, and NPQ increased in both leaf sides, but to a much higher extent at the lower compared to the upper leaf side. q_N was the best indicator for non-photochemical quenching even during a stronger HL-stress, whereas q_CN and NPQ decreased with progressive stress even though non-photochemical quenching still continued. It is strongly recommended to determine, in addition to the classical fluorescence parameters, via the PAM fluorometer also the Chl fluorescence decrease ratio R_Fd (F_d/F_s), which, when measured at saturation irradiance is directly correlated to the net CO₂ assimilation rate (_{P N}) of leaves. This R_Fd-ratio can be determined from the Chl fluorescence induction kinetics measured with the PAM fluorometer using continuous saturating light (cSL) during 4–5 min. As the R_Fd-values are fast measurable indicators correlating with the photosynthetic activity of whole leaves, they should always be determined via the PAM fluorometer parallel to the other Chl fluorescence coefficients and ratios.     

Photosynthetic Activity in Seed Wings of Dipterocarpaceae in a Masting Year: Does Wing Photosynthesis Contribute to Reproduction?

The photosynthetic rate of seed wings developed from sepals was compared with the leaf photosynthetic rate in nine dipterocarp tree species (Dipterocarpus pachyphyllus, Dryobalanops aromatica, Dryobalanops lanceolata, Shorea beccariana, Shorea ferruginea, Shorea macroptera ssp. bailonii, Shorea macroptera ssp. macropterifolia, Shorea pilosa, and Vatica spp.). The wings showed positive photosynthetic activity, but at much lower rates than in the leaves. The daily CO₂ uptake of wings showed slightly negative values in diurnal gas exchange measurements, even in D. aromatica that showed the highest photosynthetic capacity of all nine species. This low photosynthetic rate in the wings may be the result of low nitrogen and chlorophyll contents in the wing compared with leaves. However, the wings had a higher C/N ratio than leaves, and were thicker. Hence, dipterocarp wings have physical strength and defence against herbivores as higher priorities than photosynthetic activity.     

A Test of Gas Exchange Measurements on Excised Canopy Branches of Ten Tropical Tree Species

We studied gas exchange of leaves on branches that had been cut and then re-cut under water to assess the utility of measuring gas exchange on leaves of excised canopy branches. There was large variation between species in their ability to photosynthesize following excision. Some species maintained up to 86.5% of intact photosynthetic rate 60 min after excision, whereas other species dropped below 40% of intact photosynthetic rates within 3 min. Three species showed significant reductions in maximum rates of gross photosynthetic rate (P _G) on leaves of excised branches relative to intact branches. Excision significantly reduced carboxylation rates (Vc _max) in four species and electron transport (J _max) in two species. There were also significant increases in compensation irradiance and reductions of day rates of respiration relative to intact measurements. While gas exchange on excised branches can provide useful measures for canopy species, responses of individual species to branch excision need to be taken into account. Measurements on pre-screened species allow a greater understanding of canopy photosynthesis of large trees when canopy access is not an option.     

Characteristics of Photosynthetic Apparatus in Mn-Starved Maize Leaves

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

Chlorophyll fluorescence imaging of photosynthetic activity in sun and shade leaves of trees

The differences in pigment levels, photosynthetic activity and the chlorophyll fluorescence decrease ratio R _Fd (as indicator of photosynthetic rates) of green sun and shade leaves of three broadleaf trees (Platanus acerifolia Willd., Populus alba L., Tilia cordata Mill.) were compared. Sun leaves were characterized by higher levels of total chlorophylls a + b and total carotenoids x + c as well as higher values for the weight ratio chlorophyll (Chl) a/b (sun leaves 3.23–3.45; shade leaves: 2.74–2.81), and lower values for the ratio chlorophylls to carotenoids (a + b)/(x + c) (with 4.44–4.70 in sun leaves and 5.04–5.72 in shade leaves). Sun leaves exhibited higher photosynthetic rates P _N on a leaf area basis (mean of 9.1–10.1 μmol CO₂ m⁻² s⁻¹) and Chl basis, which correlated well with the higher values of stomatal conductance G _s (range 105–180 mmol m⁻² s⁻¹), as compared to shade leaves (G _s range 25–77 mmol m⁻² s⁻¹; P _N: 3.2–3.7 μmol CO₂ m⁻² s⁻¹). The higher photosynthetic rates could also be detected via imaging the Chl fluorescence decrease ratio R _Fd, which possessed higher values in sun leaves (2.8–3.0) as compared to shade leaves (1.4–1.8). In addition, via R _Fd images it was shown that the photosynthetic activity of the leaves of all trees exhibits a large heterogeneity across the leaf area, and in general to a higher extent in sun leaves than in shade leaves.     

Diurnal variation of gas exchange, chlorophyll fluorescence, and xanthophyll cycle components of maize hybrids released in different years

Diurnal variation of gas exchange, chlorophyll (Chl) fluorescence, and xanthophyll cycle components of three maize (Zea mays L.) hybrids released in different years, i.e. Baimaya (1950s), Zhongdan2 (1970s), and Nongda108 (1990s), were compared. On cloudless days, the newer hybrids always had higher net photosynthetic rate (P _N), especially at noon, than the older ones. At noon, all the hybrids decreased their maximal yield of photosystem 2 (PS2) photochemistry (F_v/F_m) and actual quantum yield of PS2 (Φ_PS2), the newer ones always showing higher values. Generally, the newer hybrids displayed higher photochemical quenching of Chl (q_P) and lower non-photochemical quenching (NPQ). The interhybrid differences in P _N may be owing to their differential photochemical efficiency. A midday depression in P _N occurred in all hybrids, which might be caused by serious photoinhibition or by decreased stomatal conductance. However, midday depression in P _N was more obvious in the older hybrids, especially when leaves were senescent. The higher de-epoxidation state of the xanthophylls was noted in older hybrids, which was confirmed by their larger NPQ. The newer maize hybrids did not need a strong de-epoxidation state since they had a better photosynthetic quantum conversion rate and a lower NPQ.     

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

The photoprotection of energy dissipation and water-water cycle were investigated by comparing chilling sensitivity of photosystems 2 (PS2) and 1 (PS1) in two chilling-sensitive plants, cucumber and sweet pepper, upon exposure to 4 °C under low irradiance (100 μmol m⁻² s⁻¹) for 6 h. During chilling stress, the maximum photochemical efficiency of PS2 (F_v/F_m) decreased only slightly in both plants, but the oxidisable P700 decreased markedly, which indicated that PS1 was more sensitive to chilling treatment under low irradiance than PS2. Sweet pepper leaves had lower F_v/F_m, higher non-photochemical quenching (NPQ), and higher oxidisable P700 during chilling stress. Activity of superoxide dismutase (SOD) and ascorbate peroxidase (APX) in cucumber leaves was higher, but APX activity decreased apparently compared to that at room temperature. The productions of active oxygen species (H₂O₂, O₂ ⁻) increased in both plants, faster in cucumber leaves than in sweet pepper leaves. In sweet pepper leaves, a stronger de-epoxidation of the xanthophyll cycle pigments, a higher NPQ could act as a major protective mechanism to reduce the formation of active oxygen species during stress. Thus sensitivity of both plants to chilling under low irradiance was dominated by the protective mechanisms between PS1 and PS2, especially the energy dissipation and the water-water cycle. This revised version was published online in August 2006 with corrections to the Cover Date.     

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.     

Seasonal changes in photosynthetic characteristics of Pachysandra terminalis (Buxaceae), an evergreen woodland chamaephyte,in the cool temperate regions of Japan

Summary Seasonal changes in photosynthetic capacity, and photosynthetic responses to intercellular CO₂ concentration and irradiance were investigated under laboratory conditions on intact leaves of Pachysandra terminalis. Photosynthetic capacity and stomatal conductance under saturating light intensity and constant water vapor pressure deficit showed almost the same seasonal trend. They increased from early June just after the expansion of leaves, reached the maximum in late-Septemer, and then decreased to winter. In over-wintering leaves they recovered and increased immediately after snow-melting, reached a first maximum in late April, and then decreased to early July in response to the reduction of light intensity on the forest floor. There-after, they increased from mid August, reached a second maximum in late September, and then decreased to winter. The parallel changes of photosynthesis and stomatal conductane indicate a more or less constant intercellular CO₂ concentration throughout the year. The calculated values of relative stomatal limitation of photosynthesis were nearly constant throughout the year, irrespective of leaf age. The results indicate that the seasonal changes in light-saturated photosynthetic capacity are not due to a change of stomatal conductance, but to a change in the photosynthetic capacity of mesophyll. Indeed, carboxylation efficiency assessed by the inital slope of the Ci-photosynthesis curve changed in proportion to seasonal changes of the photosynthetic capacity in both current-year and over-wintered leaves. High photosynthetic capacity in current-year leaves as compared with one-year-old leaves was also due to the high photosynthetic capacity of mesophyll. Nevertheless, stomatal conductance changed in proportion to photosynthetic capacity, indicating that stomatal conductance is regulated by the mesophyll photosynthetic capacity such that the intercellular CO₂ concentrations are maintained constant. The quantum yield also changed seasonally parallel with that in the photosynthetic capacity.Contribution No. 2893 from the Institute of Low Temperature Science     

Photosystem 2 Efficiency and Thylakoid Protein Pattern in DCMU-Treated Wheat Seedlings during Senescence

Changes in various components of photosynthetic apparatus during the 6-d dark incubation at 25 °C of detached control and DCMU-treated Triticum aestivum L. leaves were examined. The rate of photosystem 2 (PS2) activity was decreased with increase of the time of dark incubation in control leaves. In contrast to this, DCMU-treated leaves demonstrated high stability by slowing down the inactivation processes. Diphenyl carbazide and NH₂OH restored the PS2 activity more in control leaves than in DCMU-treated leaves. Mn²⁺ failed to restore the PS2 activity in both control and DCMU-treated samples. Similar results were obtained when F_v/F_m was evaluated by chlorophyll fluorescence measurements. The marked loss of PS2 activity in dark incubated control leaves was primarily due to the loss of D1, 33, and 23 kDa extrinsic polypeptides and 28-25 kDa LHCP2 polypeptides.