The effects of elevated light on Photosystem II function: A thermoluminescence study

We have used the technique of thermoluminescence (TL) to investigate high-light-induced chlorophyll fluorescence quenching phenomena in barley leaves, and have shown it to be a powerful tool in such investigations. TL measurements were taken from wild-type and chlorina f2 barley leaves which had been dark-adapted or exposed to 20 min illumination of varying irradiance or given varying periods of recovery following strong irradiance. We have found strong evidence that there is a sustained trans-thylakoid pH in leaves following illumination, and that this pH gives rise to quenching of chlorophyll fluorescence which has previously been identified as a slowly-relaxing component of antenna-related protective energy dissipation; we have identified a state of the PS II reaction centre resulting from high light treatments which is apparently able to perform normal charge separation and electron transport but which is non-photochemically quenched, in that the application of a light pulse of high irradiance cannot cause the formation of a high fluorescent state; and we have provided evidence that a transient state of the PS II reaction centre is formed during recovery from such high light treatments, in which electron transport from Q_Ato Q_Bis apparently impaired.     

The influence of chlorophyll fluorescence on the light gradients and the phytochrome state in a green model leaf under natural conditions

Abstract. The effect of chlorophyll fluoresence on the spectral light gradients within a model green leaf was examined under different light qualities (day-light, sunset, canopy) and different quantum efficiencies. Light fluxes within the leaf tissue are nearly doubled in the emission domain of fluorescence but the effect on the phytochrome photoequilibrium is very small.     

田间条件下棉花幼叶光合特性及光保护机制

通过比较棉花(Gossypium hirsutum)幼叶和完全展开叶气体交换参数及叶绿素荧光特性的差异, 探讨高光强下幼叶的光抑制程度及明确光保护机制间的协调机理。在田间自然条件下, 以棉花刚展平的幼嫩叶片(幼叶)和面积已达到最大的完全展开叶片为研究对象, 通过测定不同发育阶段叶片气体交换参数及叶绿素a荧光参数的变化, 并运用Dual-PAM100对不同发育阶段的叶片进行快速光响应曲线的拟合。结果表明: 幼叶和完全展开叶片在光合、荧光特性方面表现出明显的差异。与完全展开叶相比, 较低的叶绿素(Chl)含量和气孔导度(G_s)是幼叶较低净光合速率(P_n)的限制因素, 从而直接导致其光系统II (PSII)实际光化学效率(Φ_PSII)和光化学猝灭系数(q_P)的降低。在1800 μmol·m^-2·s^-1光强以下, 完全展开叶具有较强的围绕PSI循环的电子流(CEF), 有利于合成ATP, 是其具有较高光合能力的原因之一。相同光强下, 幼叶较低的光饱和点(LSP)更易受光抑制, 但其PSII原初光化学效率(F_v/F_m)的日变化幅度显著小于完全展开叶, 说明强光下幼叶通过类胡萝卜素(Car)猝灭单线态氧、光呼吸(P_r)、热耗散(NPQ)以及PSI-CEF等光保护机制能有效地耗散过剩的光能, 从而避免其光合机构发生光抑制。     

鸢尾(Iris L.)叶片取向与其光合特性及光抑制的关系

通过气体交换、叶绿素荧光、反射光谱等方法,研究了鸢尾叶片取向对植株光合特性及光抑制的影响.自然状态下,鸢尾的叶片不同取向影响植株对光能的截获;叶片净光合速率Pn与光合有效辐射PAR呈极显著相关;东西取向叶片的Pn要大于南北取向.南北取向的植株中叶片叶绿素(Chl a和Chl b),类胡萝卜素(Car)含量略高于东西取向.日进程中,各取向的叶片在一天中均没有发生明显的光抑制.相对于东西取向的植株,南北取向植株发生了明显的倾斜;在两种取向的植株中,叶片东侧和南侧的光化学反射指数(PRI)下调幅度较大;PRI的变化量(△PRI)大小依次为:东侧>南侧>西侧>北侧.鸢尾植株取向改变了叶片倾斜角度,两者共同导致光能截获减小;同时,叶片光能利用效率下调和叶黄素循环增强,这可能是不同取向植株均未发生严重光抑制的原因.     

叶角、光呼吸和热耗散协同作用减轻大豆幼叶光抑制

研究了大豆叶片逐步展开过程中的色素组成、气体交换、荧光动力学以及叶片角度等特性。随着叶片展开程度的增加 ,叶绿素含量和叶绿素 a/ b比值增加 ;光合速率 (Pn)也增加 ,揭示叶片展开过程中光合机构是逐步完善的。自然状态下 ,不同展开程度的叶片均未发生明显的光抑制 ;但将叶片平展并暴露在 12 0 0μmol/ (m2 · s)光下时幼叶发生严重的光抑制 ,伴随叶面积的增加光抑制程度减轻。强光下 ,尽管幼叶光呼吸 (Pr)的测定值较低 ,但幼叶光呼吸与总光合之比 (Pr/ Pm)较高。将叶片平展置于强光下时 ,幼叶的实际光化学效率 (ΦPSII)明显下调 ,非光化学猝灭 (NPQ)大幅增加 ;幼叶叶黄素库较大 ,光下积累较多的脱环氧化组分 ,揭示幼叶依赖叶黄素循环的热耗散增强。自然条件下测量叶片角度 ,观察到在叶片展开过程中叶柄夹角逐渐增加 ;日动态过程中幼叶的悬挂角随光强增加而明显减小 ,完全展开叶的悬挂角变化幅度很小。叶片角度的变化使实际照射到幼叶叶表的光强减少。推测较强的光呼吸、依赖叶黄素循环的热耗散以及较大的叶角变化可能是自然状态下幼叶未发生严重光抑制的原因     

干旱胁迫对沙冬青叶片防御光破坏机制的影响

沙冬青(Ammopiptanthus mongolicus (Maxim.) Cheng f.)是生长在沙漠及干旱荒漠地区的常绿灌木.在夏季,其叶片经常遭受中午强光(超过1 500 μmol*m-2*s-1) 胁迫,出现明显的光抑制现象.我们利用便携式光合测定系统(CIRAS-1)和脉冲调制荧光仪(MFMS-2)测定了自然形成的干旱胁迫条件下沙冬青光合和荧光参数的日变化,主要探讨了干旱胁迫对沙冬青叶片防御强光破坏机制的影响.结果表明,正常水分和干旱胁迫下,沙冬青叶片的净光合速率(Pn)、 PSⅡ最大光化学效率(Fv/Fm)和PSⅡ非环式电子传递效率(ΦPSⅡ)在中午都明显降低;相对正常水分条件而言,干旱胁迫下初始荧光(Fo)先下降后上升,荧光的非光化学淬灭(NPQ)上升较快并在一定水平上维持不变.由此推断晴天中午沙冬青叶片在正常水分条件下主要采取依赖叶黄素循环的热耗散机制;而在干旱胁迫条件下主要采取了依赖叶黄素循环的热耗散和PSⅡ反应中心可逆失活两种保护机制.     

Underestimate of PS2 efficiency in the field due to high leaf temperature resulting from leaf clipping and its amendment

Chlorophyll fluorescence parameter F_v/F_m, an indicator of the maximum efficiency of PS2, is routinely measured in the field with plant leaves darkened by leaf clips. I found that on a sunny day of subtropical summer, the F_v/F_m ratio was often underestimated because of a large F₀ value resulted from a high leaf temperature caused by clipping the leaf under high irradiance, especially for long (e.g. 20 min) duration. This phenomenon may overestimate the down-regulation of PS2 efficiency under high irradiance. When leaf temperature was lower than 40 °C, the F₀ level of rice leaves under clipping remained practically unchanged. However, F₀ increased drastically with leaf temperature rising over 40 °C. In most measurements, no significant difference in F_m was found between rice leaves dark-adapted by leaf clips for 10 min and for 20 min. Therefore, shading leaf clips to prevent a drastic increase of leaf temperature, using F₀ measured immediately after the leaf being darkened to calculate F_v/F_m, as well as shortening the duration of leaf clipping are useful means to avoid an underestimate of F_v/F_m.     

Ultra-structural and functional changes in the chloroplasts of detached barley leaves senescing under dark and light conditions

Changes in the chloroplast ultra-structure and photochemical function were studied in detached barley (Hordeum vulgare L. cv. Akcent) leaf segments senescing in darkness or in continuous white light of moderate intensity (90 mumol m-2 s-1) for 5 days. A rate of senescence-induced chlorophyll degradation was similar in the dark- and light-senescing segments. The Chl a/b ratio was almost unchanged in the dark-senescing segments, whereas in the light-senescing segments an increase in this ratio was observed indicating a preferential degradation of light-harvesting complexes of photosystem II. A higher level of thylakoid disorganisation (especially of granal membranes) and a very high lipid peroxidation were observed in the light-senescing segments. In spite of these findings, both the maximal and actual photochemical quantum yields of the photosystem II were highly maintained in comparison with the dark-senescing segments.     

The scaling of leaf area and mass: the cost of light interception increases with leaf size

For leaves, the light-capturing surface area per unit dry mass investment (specific leaf area, SLA) is a key trait from physiological, ecological and biophysical perspectives. To address whether SLA declines with leaf size, as hypothesized due to increasing costs of support in larger leaves, we compiled data on intraspecific variation in leaf dry mass (LM) and leaf surface area (LA) for 6334 leaves of 157 species. We used the power function LM=alpha LAbeta to test whether, within each species, large leaves deploy less surface area per unit dry mass than small leaves. Comparing scaling exponents (beta) showed that more species had a statistically significant decrease in SLA as leaf size increased (61) than the opposite (7) and the average beta was significantly greater than 1 (betamean=1.10, 95% CI 1.08-1.13). However, scaling exponents varied markedly from the few species that decreased to the many that increased SLA disproportionately fast as leaf size increased. This variation was unrelated to growth form, ecosystem of origin or climate. The average within-species tendency found here (allometric decrease of SLA with leaf size, averaging 13%) is in accord with concurrent findings on global-scale trends among species, although the substantial scatter around the central tendency suggests that the leaf size dependency does not obligately shape SLA. Nonetheless, the generally greater mass per unit leaf area of larger than smaller leaves directly translates into a greater cost to build and maintain a unit of leaf area, which, all else being equal, should constrain the maximum leaf size displayed.     

Interactive effects of water,light and heat stress on photosynthesis in Fremont cottonwood

Fremont cottonwood seedlings are vulnerable to water stress from rapid water‐table decline during river recession in spring. Water stress is usually cited as the reason for reduced establishment, but interactions of water stress with microclimate extremes are more likely the causes of mortality. We assessed photosynthetic responses of Fremont cottonwood seedlings to water, light and heat stresses, which commonly co‐occur in habitats where seedlings establish. Under moderate temperature and light conditions, water stress did not affect photosynthetic function. However, stomatal closure during water stress predisposed Fremont cottonwood leaves to light and heat stress, resulting in greatly reduced photosynthesis beginning at 31 °C versus at 41 °C for well‐watered plants. Ontogenetic shifts in leaf orientation from horizontal to vertical, which occur as seedlings mature, reduce heat and light stress, especially during water stress. When compared with naturally occurring microclimate extremes, seedling stress responses suggest that reduced assimilation and photoprotection are common for Fremont cottonwood seedlings on exposed point bars where they establish. These reductions in photosynthesis likely have negative impacts on growth and may predispose young (<90‐day‐old) seedlings to early mortality during rapid water‐table declines. Interactions with heat and light stress are more important in these effects than water stress alone.     

Acclimation of tropical tree seedlings to excessive light in simulated tree-fall gaps

Acclimation to periodic high‐light stress was studied in tree seedlings from a neotropical forest. Seedlings of several pioneer and late‐succession species were cultivated under simulated tree‐fall gap conditions; they were placed under frames covered with shade cloth with apertures of different widths that permitted defined periods of daily leaf exposure to direct sunlight. During direct sun exposure, all plants exhibited a marked reversible decline in potential photosystem II (PSII) efficiency, determined by means of the ratio of variable to maximum Chl a fluorescence (F_v/F_m). The decline in F_v/F_m under full sunlight was much stronger in late‐succession than in pioneer species. For each gap size, all species exhibited a similar degree of de‐epoxidation of violaxanthin in direct sunlight and similar pool sizes of xanthophyll cycle pigments. Pool sizes increased with increasing gap size. Pioneer plants possessed high levels of β‐carotene that also increased with gap size, whereas α‐carotene decreased. In contrast to late‐succession plants, pioneer plants were capable of adjusting their Chl a/b ratio to a high value in wide gaps. The content of extractable UV‐B‐absorbing compounds was highest in the plants acclimated to large gaps and did not depend on the successional status of the plants. The results demonstrate a better performance of pioneer species under high‐light conditions as compared with late‐succession plants, manifested by reduced photoinhibition of PSII in pioneer species. This was not related to increased pool size and turnover of xanthophyll cycle pigments, nor to higher contents of UV‐B‐absorbing substances. High β‐carotene levels and increased Chl a/b ratios, i.e. reduced size of the Chl a and b binding antennae, may contribute to photoprotection in pioneer species.     

Changes in photosynthesis,xanthophyll cycle,and sugar accumulation in two North Australia tropical species differing in leaf angles

Two tropical species of North Australia, Acacia crassicarpa and Eucalyptus pellita, have similar leaf size and leaf structure but different leaf angles. A. crassicarpa with near vertical leaf angle directly reduced photon absorption and leaf temperature (T _l) and had relatively high photosynthetic activity (P _max) and low xanthophyll cycle activity. In contrast, E. pellita with a small leaf angle exhibited high T _l, low P _max, and high activity of xanthophyll cycle which was useful for the dissipation of excessive energy and reduction of photoinhibition. In the dry season, contents of soluble sugars including pinitol, sucrose, fructose, and glucose in A. crassicarpa increased whereas larger amounts of only fructose and glucose were accumulated in E. pellita. Different sugar accumulation may be involved in osmotic adjustment of leaves during water stress that makes photosynthesis more efficient. The leaf angle may be critical for developing different protective mechanisms in these two tropical tree species that ensure optimal growth in the high irradiance and drought stress environment in North Australia.     

Susceptibility of CAM Dendrobium leaves and flowers to high light and high temperature under natural tropical conditions

Responses of CAM Dendrobium Sonia leaves and flowers to high light and high temperature were studied in shade-grown plants after exposure to intermediate and full sunlight under natural conditions. Photosynthetic O₂ evolution decreased in leaves after exposure to full sunlight for 2 weeks while leaves exposed to intermediate sunlight showed an increase in photosynthesis as compared to those leaves maintained in the shade. On the first day of treatment, the changes of F_v/F_m in both leaves and petals grown in the shade were negligible during the day. However, there was a steep decrease in F_v/F_m in both leaves and petals with an increase in incident light during midday after exposure to full sunlight. When exposed to intermediate sunlight, there were no significant changes in F_v/F_m in leaves. The F_v/F_m values of petals, however, decreased during midday. Temperature of thin petals was higher than that of thick leaves during midday under full and intermediate sunlight while that of petals and leaves were similar when grown in the shade. Over the 2-week treatment period, lowered chlorophyll and sustained decreases in F_v/F_m were observed in both leaves and flowers (sepals and petals) when exposed to full sunlight, indicative of ‘chronic photoinhibition'. Photoinhibition was prevented in leaves but occurred in flowers when exposed to intermediate sunlight. It was assumed that photodamage to both leaves and flowers were partially due to the higher temperature. The higher susceptibility of flowers to high light as compared to that of leaves was due to its higher temperature during midday. This was further supported by the findings that more severe damage occurred in flowers at higher temperature of 38°C than 28°C under a higher PFD of 1500 μmol m⁻² s⁻¹.     

Influence of irradiance,dissolved oxygen concentration,and temperature on the growth of <Emphasis Type="Italic">Chlorella sorokiniana</Emphasis>

The growth response of Chlorella sorokiniana to certain irradiance, DO, and temperature demonstrated the possible causes of low productivity with this strain in outdoor cultures. The growth (biomass productivity) and chlorophyll fluorescence (F_v/F_m) were substantially reduced when the dissolved oxygen (above 200 % of air saturation) and temperature were elevated.     

Susceptibility to photoinhibition of three deciduous broadleaf tree species with different successional traits raised under various light regimes

The susceptibility to photoinhibition of tree species from three different successional stages were examined using chlorophyll fluorescence and gas exchange techniques. The three deciduous broadleaf tree species were Betula platyphylla var. japonica, pioneer and early successional, Quercus mongolica, intermediate shade‐tolerant and mid‐successional, and Acer mono, shade‐tolerant and late successional. Tree seedlings were raised under three light regimes: full sunlight (open), 10% full sun, and 5% full sun. Susceptibility to photoinhibition was assessed on the basis of the recovery kinetics of the ratio of vaviable to maximum fluorescence (F_v/F_m) of detached leaf discs exposed to about 2000 μmol m^?1 s^?1 photon flux density (PFD) for 2 h under controlled conditions (25 to 28 °C, fully hydrated). Differences in susceptibility to photodamage among species were not significant in the open and 10% full sun treatments. But in 5% full sun, B. platyphylla sustained a significantly greater photodamage than other species, probably associated with having the lowest photosynthetic capacity indicated by light‐saturated photosynthetic rate (B. platyphylla, 9·87, 5·85 and 2·82; Q. mongolica, 8·05, 6·28 and 4·41; A. mono, 7·93, 6·11 and 5·08 μmol CO₂ m^?1 s^?1for open, 10% and 5% full sun, respectively). To simulate a gap formation and assess its complex effects including high temperature and water stress in addition to strong light on the susceptibility to photoinhibition, we examined photoinhibition in the field by means of monitoring ΔF/F′_m on the first day of transfer to natural daylight. Compared with ΔF/F′_m in AM, the lower ΔF/F′_m in PM responding to lower PFD following high PFD around noon indicated that photoinhibition occurred in plants grown in 10 and 5% full sun. The diurnal changes of ΔF/F′_m showed that Q. mongolica grown in 5% full sun was less susceptible to photoinhibition than A. mono although they showed little differences both in photosynthetic capacity in intact leaves and susceptibility to photoinhibition based on leaf disc measurements. These results suggest that shade‐grown Q. mongolica had a higher tolerance for additional stresses such as high temperature and water stress in the field, possibly due to their lower plasticity in leaf anatomy to low light environment.     

Dependence of photosynthesis and energy dissipation activity upon growth form and light environment during the winter

Two very distinctive responses of photosynthesis to winter conditions have been identified. Mesophytic species that continue to exhibit growth during the winter typically exhibit higher maximal rates of photosynthesis during the winter or when grown at lower temperatures compared to individuals examined during the summer or when grown at warmer temperatures. In contrast, sclerophytic evergreen species growing in sun-exposed sites typically exhibit lower maximal rates of photosynthesis in the winter compared to the summer. On the other hand, shaded individuals of those same sclerophytic evergreen species exhibit similar or higher maximal rates of photosynthesis in the winter compared to the summer. Employment of the xanthophyll cycle in photoprotective energy dissipation exhibits similar characteristics in the two groups of plants (mesophytes and shade leaves of sclerophytic evergreens) that exhibit upregulation of photosynthesis during the winter. In both, zeaxanthin + antheraxanthin (Z + A) are retained and PS II remains primed for energy dissipation only on nights with subfreezing temperatures, and this becomes rapidly reversed upon exposure to increased temperatures. In contrast, Z + A are retained and PS II remains primed for energy dissipation over prolonged periods during the winter in sun leaves of sclerophytic evergreen species, and requires days of warming to become fully reversed. The rapid disengagement of this energy dissipation process in the mesophytes and shade sclerophytes apparently permits a rapid return to efficient photosynthesis and increased activity on warmer days during the winter. This may be associated with a decreasing opportunity for photosynthesis in source leaves relative to the demand for photosynthesis in the plant's sinks. In contrast, the sun-exposed sclerophytes – with a relatively high source to sink ratio – maintain PS II in a state primed for high levels of energy dissipation activity throughout much of the winter. Independent of whether photosynthesis was up- or downregulated, all species under all conditions exhibited higher levels of soluble carbohydrates during the winter compared to the summer. Thus downregulation of photosynthesis and of Photosystem II do not appear to limit carbohydrate accumulation under winter conditions. A possible signal communicating an altered source/sink balance, or that may be influencing the engagement of Z + A in energy dissipation, is phosphorylation of thylakoid proteins such as D1.This revised version was published online in October 2005 with corrections to the Cover Date.     

Light adaptation of the phytoplankton diatom Phaeodactylum tricornutum under conditions of natural light climate

Cells of Phaeodactylum tricornutum were precultured under axenic conditions in a full medium and then exposed to natural light conditions at various depths in the eutrophic lake „Meerfelder Maar”︁ (Eifel, FRG) for several days. After exposition the cells were characterized with respect to growth parameters, photosynthetic performance and xanthophyll cycle pigments. In order to test the resistance of the cells grown at different depths against photostress, the cells were illuminated with photoinhibitory light. The variable chlorophyll a-fluorescence and the oxygen quantum yield at a non-saturating light intensity were simultaneously measured after photostress and subsequent recovery. The xanthophyll cycle pigments and the content in α-tocopherol were monitored during photostress to get molecular information about the physiological reasons of light-stress resistance. The data give evidence that cells grown close to the surface show a faster decline in photosynthetic performance and a more efficient recovery than cells from lower depths. There is clear indication that under natural conditions when the light is fluctuating between optimal, sub- and supraoptimal intensities the photostress resistance is much higher than under conditions of the absence of light stress. The molecular basis for light stress resistance seems to be the pool size and the conversion kinetics of the xanthophyll cycle pigments and the capacity of the oxygen-scavenging system. The effect of in-situ light adaptation is discussed with respect to the assessment of the potential of the primary production.     

黄瓜幼苗子叶在低温下的光抑制及其恢复

用PAM脉冲调制荧光仪测定叶绿素荧光的变化研究了黄瓜幼苗子叶在PFD为50和100μmolm-2s-1,温度为4、7、10、15℃下的光抑制及其恢复。结果表明,黄瓜幼苗子叶Fv／Fm随着温度的下降和PFD的增加而下降,并且增加等量的PFD在4℃下比在10℃下引起更大的Fv／Fm下降。在黑暗条件下光抑制有轻微恢复,但完士恢复必需光照,且恢复起始时的光照十分重要。DTT可部分抑制叶绿素荧光Fo和Fm的猝灭,且15℃下比在4℃下抑制效果更大。CAP能强烈地加剧光抑制并几乎完全抑制恢复,且10℃下比在4℃下对光抑制的加剧作用更大。冷锻炼提高了黄瓜幼苗抵抗低温先抑制的能力,而CAP对冷锻炼苗比未锻炼苗的低温光抑制具有更大的加剧作用。     

The consequences of delayed greening during leaf development for light absorption and light use efficiency

Many species of rainforest plants have an unusual form of leaf development such that leaves delay greening until after full leaf expansion. Chlorophyll accumulation was measured during leaf development in five woody rainforest species, three with white young leaves, and two with ‘normal’ greening. In the three species with white leaves, the chlorophyll content of the expanding leaves was about 0.4mg dm^?2, whereas in the two species with green young leaves, chlorophyll content was about 2.1 mg dm^?2. Chlorophyll accumulation in greenhouse and field experiments was independent of light level. During leaf expansion, species with delayed chloroplast development only absorb 18–25% of the maximum possible light, compared with 80% for species with normal greening. Furthermore, species with delayed greening have low chlorophyll contents and reduced absorption for at least 30 d after full expansion. At a PPFD typical of the forest under story, the photosynthetic light use efficiency based upon incident radiation was 0.030–0.036 for species with delayed chloroplast development and 0.068–0.085 for the two species with normal greening. The lower light use efficiency of white species was primarily due to decreased light absorption. However, they also had a slightly lower light use efficiency based upon absorbed radiation, suggesting that development of other components of the photo-synthetic apparatus also may be delayed. Despite the fact that delayed greening decreases light absorption and light use efficiency during leaf development, it is extremely common in shade-tolerant species. We suggest that an advantage of delayed greening is that resources are not invested in the leaf until it is fully expanded and better defended from herbivores.