C_3植物光合效率的日变化

多种田间C_3植物在晴天的光合效率常有明显的日变化,中午前后光合效率降低。C_3植物大豆叶片光合效率中午降低的主要原因,不是空气CO_2浓度和气孔导度及光呼吸的变化,而可能是光抑制。因为:1.在饱和CO_2中测定仍可观测到这种中午降低;2.光合作用的饱和光强远低于晴天中午的太阳光强;3.用纱布预遮阴可以提高叶片的光合效率;4.阴天时叶片光合效率不发生中午降低。     

光合作用对光和二氧化碳响应的观测方法探讨

用便携式光合仪LI-6400观测自然条件下生长的盆栽蚕豆叶片光合作用对光和二氧化碳的响应发现：（1）用未经过光合诱导的叶片观测光合作用对光的响应会得到即使在全太阳光强下光合作用仍然不饱和的假象;（2）利用某些经验方程计算的饱和光强远低于实际观测值;（3）在观测光合作用对CO2的响应过程中,每一次CO2浓度变化都应当伴随一次光合仪的匹配步骤,否则所得结果偏差很大;（4）在不饱和光下观测光合作用对CO2的响应,会导致对叶片光合能力的低估。     

Two patterns of leaf photosynthetic response to irradiance transition from saturating to limiting one in some plant species

Plants often regulate the amount and size of light-harvesting antenna (LHCII) to maximize photosynthesis at low light and avoid photodamage at high light. Gas exchange, 77 K chlorophyll fluorescence, photosystem II (PSII) electron transport as well as LHCII protein were measured in leaves irradiated at different light intensities. After irradiance transition from saturating to limiting one leaf photosynthetic rate in some species such as soybean and rice declined first to a low level, then increased slowly to a stable value (V pattern), while in other species such as wheat and pumpkin it dropped immediately to a stable value (L pattern). Saturating pre-irradiation led to significant declines of both 77 K fluorescence parameter F685/F735 and light-limited PSII electron transport rate in soybean but not in wheat leaves, indicating that some LHCIIs dissociate from PSII in soybean but not in wheat leaves. The L pattern of LHCII-decreased rice mutant and the V pattern of its wild type demonstrate that the V pattern is linked to dissociation/reassociation of some LHCIIs from/to PSII.     

不同氮营养水平下草莓叶片光合作用对高CO2浓度的适应

研究了不同氮素水平（12mmol／L,4mmol／L,0、4mmol／L）下生长的‘丰香’草莓在富C02（700μL/L）和大气CO（390μL/L）下的光合作用。结果表明,高氮（12mmol／L）下,在富CO2环境中生长的‘丰香’草莓叶片未出现光合作用下调,富CO2下草莓叶片的净光合速率、最大羧化速率（Vc.max）、最大电子传递速率（Jmax）、碳同化的电子传递速率（Jc）和光化学猝灭系数（qp）等均显著提高;而在中氮（4mmol／L）、低氮（0．4mmol／L）下,富CO2下生长的草莓叶片的上述参数均出现不同程度的下降。富CO2下,无论氮素水平如何,草莓叶片的光呼吸电子传递速率（Jo）均降低高氮草莓叶片的非光化学猝灭系数（qN或NPQ）降低,光抑制降低,而低氮则相反。上述结果说明,氮素供应不足时草莓叶片在富CO2下光合作用出现下调,因此生产上进行CO2施肥时应适度增加氮素的供应。     

珊瑚树阳生和阴生叶片光合特性和状态转换的比较

珊瑚树阳生和阴生叶片是在不同光照环境中长期生长的,它们的光合特性有一些明显的差异.与阳生叶片相比,阴生叶片单位干重的叶绿素含量较多,类囊体膜垛叠程度较高(即每个基粒的类囊体膜垛叠层数较多,基粒类囊体的直径较大),而叶绿素a/b比值、光合作用的饱和光强和最大净光合速率等较低.用弱红光诱导阳生和阴生叶片向状态2转换时,叶绿素荧光Fm/Fo和F685/F735先迅速下降再逐渐回升,这表明两种叶片都先后通过满溢和LHCⅡ转移调节激发能在PSⅡ和PSⅠ之间的分配,改善光能利用,但阳生叶片Fm/Fo和F685/F735下降的幅度较大.     

Photosynthetic Adaptation to Light Intensity in Plants Native to Shaded and Exposed Habitats

Photosynthetic adaptation to light intensity has been studied in clones of populations from shaded and exposed habitats of Rumex acetosa and Geum rivale. Clones of the shade species Lamium galeobdolon and the sun species Plantago lanceolata were also included for comparison. The plants were grown under controlled conditions at a high and a low light intensity. The capacity of photosynthetic carbon dioxide uptake at low as well as at saturating light intensities was determined on single attached leaves. As was previously demonstrated in Solidago virgaurea, clones of populations native to shaded and to exposed environments show differences in the photosynthetic response to light intensity during growth. The data provide evidence that populations of the same species native to habitats with contrasting light intensities differ in their photosynthetic properties in an adaptive manner Ln a similar mode as sun and shade species.     

Photosynthetic responses to variable light: a comparison of species from contrasting habitats

Photosynthetic responses to variable light were compared for species from habitats differing in light availability and dynamics. Plants were grown under the same controlled conditions and were analysed for the kinetics of photosynthetic induction when photon flux density (PFD) was increased from 25 to 800 mol m^-2s^-1. Gas exchange techniques were used to analyse the two principal components of induction, opening of stomata and activation of ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco). On average, 90% of the final photosynthetic rate was attained after 7 min for obligate shade plants (two species), 18 min for fast-growing sun plants (seven species from productive habitats) and 32 min for slow-growing sun plants (nine species from unproductive habitats). The rapidity of response of the shade plants was explained by stomata remaining more open in the low-light period prior to induction. This was also observed in two species of deciduous trees, which therefore resembled shade plants rather than other fast-growing sun plants. The slow response of the slow-growing sun plants was the result of lower rates of both Rubisco activation and stomatal opening, the latter being more important for the final phase of induction. The lower rate of Rubisco activation was confirmed by direct, enzymatic measurements of representative plants. With increasing leaf age, the rate of stomatal opening appeared to decrease but the rate of Rubisco activation was largely conserved. Representative species were also compared with respect to the efficiency of using light-flecks relative to continuously high light. The shade plants and the slow-growing sun plants had a higher efficiency than the fast-growing sun plants. This could be related to the presence of a higher electron transport capacity relative to carboxylation capacity in the former group, which seems to be associated with their lower photosynthetic capacities. Representative species were also compared with respect to the ability to maintain the various induction components through periods of low light. Generally, the fast-growing sun plants were less able than the other two categories to maintain the rapidly reversible component. Thus, although the rate of induction appears to be related to the ecology of the plant, other aspects of photosynthetic dynamics, such as the efficiency of using lightflecks and the ability to maintain the rapidly reversible component, seem rather to be inversely related to the photosynthetic capacity.     

青藏高原药用植物唐古特山莨菪和唐古特大黄光合作用对强光的响应

与唐古特大黄相比,唐古特山莨菪的表观光合量子效率(AQY)较高,但最大净光合速率(Pmax)较低。在光强小于1200μmolm-2s-1时,后者用于碳同化的电子传递占总光合电子传递的比例(JC/JF)比前者高,而分配于光呼吸的电子传递(JO/JF)及Rubisco氧化和羧化速率的比值(VO/VC)则相反;光强大于1200μmolm-2s-1以后两种植物的这些参数都趋向稳定。随光强增加,后者叶片吸收光能分配于热耗散(D)的增加斜率较前者高,表明两高山植物对强辐射的适应方式略有不同。加强光呼吸途径的耗能代谢和PSII天线热耗散份额是唐古特山莨菪适应高原强辐射的主要方式,而提高叶片光合能力则是唐古特大黄的一种适应方式。     

4种木本植物叶片的光合电子传递和吸收光能分配特性对光强的适应

以气体交换和叶绿素荧光测定相结合的方法研究了亚热带自然林乔木荷树、黧蒴和林下灌木九节、罗伞幼苗的光合电子传递及激发能利用的分配对生长光强的适应特性。4种植物生长于100%、36%和16%的自然光下8个月,叶片的光化学速率和热能耗散速率随光强增大而提高,热能耗散占总的光能吸收的比例也因光强不同而改变,16%光下的相对热耗散率约为40%～45%,100%自然光下增大至50%～75%。叶片总的非环式电子流速率及其分配到光呼吸的比例在100%光强下最高。乔木和灌木的电子传递和光能分配特性在16%光下相似,在100%光下差别较明显。除灌木种有较高的热耗散比例之外,其余的参数皆比乔木的低。结果表明乔木与灌木皆可通过提高激发能热耗散比例和提高光合电子传递向光呼吸的比例来适应于高光强条件。     

Influence of light intensity during growth on photosynthesis and activity of several key photosynthetic enzymes in a C4 plant (Zea mays)

Maize ( Zea mays L. Hybrid Sweet Corn, Royal Crest), a C₄ plant, was grown under different light regimes, after which the rate of photosynthesis and activities of several photosynthetic enzymes (per unit leaf chlorophyll) were measured at different light intensities. Plants were grown outdoors under direct sunlight or 23% of direct sunlight, and in growth chambers at photosynthetic photon flux densities of about 20% and 8% of direct sunlight. The plants grown under direct sunlight had a higher light compensation point than plants grown under lower light. At a light intensity about 25% of direct sunlight, plants from all growth regimes had a similar rate of photosynthesis. Under saturating levels of light the plants grown under direct sunlight had a substantially higher rate of photosynthesis than plants grown under the lower light regimes. The higher photosynthetic capacity in the plants grown under direct sunlight was accompanied by an increased activity of several photosynthetic enzymes and in the amount of the soluble protein in the leaf. Among five photosynthetic enzymes examined, RuBP carboxylase (EC 4.1.1.39) and pyruvate, Pi dikinase (EC 2.7.9.1) were generally just sufficient to account for rates of photosynthesis under saturating light; thus, these may be rate limiting enzymes in C₄ photosynthesis. Pyruvate, Pi dikinase and NADP-malate dehydrogenase (EC 1.1.1.82) were the only enzymes examined which were light activated and increased in activity with increasing light intensity. In the low light grown plants the activity of pyruvate, Pi dikinase closely paralleled the photosynthetic rate measured under different light levels. With the plants grown under direct sunlight, as light intensity was increased the activation of pyruvate, Pi dikinase and NADP⁺-malate dehydrogenase proceeded more rapidly than photosynthesis.     

Adaptation of the Photosynthetic Apparatus of Scenedesmus obliquus to Strong and Weak Light Conditions

Homocontinuous cultures of the unicellular green alga Scenedesmus obliquus were grown under strong (28 W/m²～28,000 lux) and weak (5 W/m²～5000 lux) light conditions to simulate the conditions of ‘sun’ and ‘shade’ plants. As in higher plants the cells adapted to strong light had less chlorophyll but demonstrated a higher photosynthetic capacity and a higher respiration rate, so that their compensation point was reached at three times higher energy than in the cells grown under low light intensities. The CO₂ fixation rate and the RuDP carboxylase activity under saturating light intensities were both higher in the cells grown in strong light. In spite of the differences in the pigment content and in the light saturated photosynthetic capacities for both cultures, the quantum yields of photosynthetic oxygen evolution were equal. As documented for some species of higher plants Scenedesmus is not genetically determined to be either a ‘sun’ or ‘shade’ organism but can adapt its photosynthetic apparatus to the different light intensities.     

不同光强下焕镛木和观光木的光合参数变化

 生长在全日光强下的焕镛木(Woonyoungia septentrionalis)和观光木(Tsoongiodendron lotungensis)幼树叶片的最大光合速率、表观量子产率和光能转换效率均较生长在40%和20%日光强的高。当生长光强从全日光强降低至40%日光强时,焕镛木的表观量子产率和光能转换效率分别降低13.1%和6.3%,而观光木则相应分别降低23.8%和33.4%。生长光强降低至40%日光强时,焕镛木的Rubisco最大羧化速率(Vcmax)未见变化;而最大电子传递速率（Jmax）则降低14.1%,表明Jmax对光强降低的响应较Vcmax敏感。当生长光强从全日光强降低到40%和20%日光强时,观光木的Vcmax分别降低7.7%和31.7%,而Jmax则分别降低9.7%和42%。光强从全日光强降低至40%日光强,焕镛木叶氮在Rubisco和捕光叶绿素蛋白复合体中的分配系数没有明显改变,而叶氮在生物力能学组分中的分配系数降低则较为明显（20.4%）,表明生长光强降低对叶氮在光合电子传递链组分分配的影响较在Rubisco的大。结果表明,焕镛木表现阳生树种特性,在迁地保育中宜选择向阳小生境种植,而观光木较耐荫,可种植在较遮荫的环境。     

Growth and photosynthetic responses of wheat plants grown in space.

Growth and photosynthesis of wheat (Triticum aestivum L. cv Super Dwarf) plants grown onboard the space shuttle Discovery for 10 d were examined. Compared to ground control plants, the shoot fresh weight of space-grown seedlings decreased by 25%. Postflight measurements of the O2 evolution/photosynthetic photon flux density response curves of leaf samples revealed that the CO2-saturated photosynthetic rate at saturating light intensities in space-grown plants declined 25% relative to the rate in ground control plants. The relative quantum yield of CO2-saturated photosynthetic O2 evolution measured at limiting light intensities was not significantly affected. In space-grown plants, the light compensation point of the leaves increased by 33%, which likely was due to an increase (27%) in leaf dark-respiration rates. Related experiments with thylakoids isolated from space-grown plants showed that the light-saturated photosynthetic electron transport rate from H2O through photosystems II and I was reduced by 28%. These results demonstrate that photosynthetic functions are affected by the microgravity environment.     

Study on photosynthetic responses and chlorophyll fluorescence in Rhizophora mucronata seedlings under shade regimes

The photosynthetic performance of mangrove Rhizophora mucronata seedlings grown under seasonally full light (HL), 50 % shade (ML), and 80 % shade (LL) conditions was characterized by gas exchange, and chlorophyll fluorescence. The carboxylation efficiency significantly affected the seasonal change of the photosynthetic capacity. Temperature and light might have synergic effect on the carboxylation efficiency. The photosynthetic rate (PN) of R. mucronata seedlings under shade regimes, however, could not be attributed to variability in chlorophyll, C _i , ΦPSII, ETR or qP values but more to differences in carboxylation efficiency, g _max, and E _max. HL and ML plants had higher PN, g _s and E than the LL ones. Nevertheless, LL leaves exhibited low photoinhibition susceptibility. The high non-photochemical quenching in HL leaves may show that applied light intensity probably exceeded the photosynthetic capability. The findings indicate that ML treatments provided the best condition to obtain such carbon fixation capacity.     

Influence of Etherel and Gibberellic Acid on Carbon Metabolism,Growth, and Essential Oil Accumulation in Spearmint (Mentha Spicata)

Controlled environment chamber and glasshouse studies were conducted on six herbaceous annual species grown at 350 (AC) and 700 (EC) mol(CO₂) mol^-1 to determine whether growth at EC resulted in acclimation of the apparent quantum yield of photosynthesis (QY) measured at limiting photosynthetic photon flux density (PPFD), or in acclimation of net photosynthetic rate (P _N) measured at saturating PPFD. It was also determined whether acclimation in P _N at limiting PPFD was correlated with acclimation of carboxylation efficiency or ribulose-1,5-bisphosphate (RuBP) regeneration rate measured at saturating PPFD. Growth at EC reduced both the QY and P _N at limiting PPFD in three of the six species. The occurrence of photosynthetic acclimation measured at a rate limiting PPFD was independent of whether photosynthetic acclimation was apparent at saturating measurement PPFD. At saturating measurement PPFD, acclimation to EC in the apparent carboxylation efficiency and RuBP regeneration capacity also occurred independently. Thus at least three components of the photosynthetic system may adjust independently when leaves are grown at EC. Estimates of photosynthetic acclimation at both high and low PPFD are necessary to accurately predict photosynthesis at the whole plant or canopy level as [CO₂] increases.     

Light-Induced Adaptive Responses under Greenhouse and Controlled Conditions in the Fern Pteris cretica var. ouvrardii

The photosynthetic capabilities of the fern Pteris cretica var. ouvrardii were analysed by means of the light response curves of CO₂ exchange. In control growth conditions (greenhouse, low-light: 20–32 W m^?2); photosynthesis was shown to be saturated for low irradiance (20–25 W m^?2); the saturating photosynthetic rate, very low as compared to higher plants, was due to an extremely high intracellular resistance. When irradiance during the photosynthesis measurement was higher than 60–80 W m^?2, a constant decline of net CO₂ exchange as a function of time was observed. When irradiance during growth was enhanced, whether in greenhouse (20–250 W m^?2) or controlled (62 W m^?2) conditions, the first fronds that had developed in the new condition from the crosier stage exhibited decreased net maximal photosynthesis and a decreased efficiency in low light, but saturating irradiance was unmodified. However, the fronds whose entire differentiation (from meristem) occurred under these moderate irradiances (plants defoliated of all fronds and crosiers at the time of transfer), possessed more efficient photosynthetic characteristics than control plants. Pteris is able to grow under extreme shade conditions (4–8 W m^?2); light saturating photosynthesis and efficiency are higher under extreme shade than under control conditions. These adaptive characteristics indicate that Pteris is a well-adapted shade species.     

午间强光胁迫下SOD对大豆叶片光合机构的保护作用

晴天田间大豆叶片Ｐｎ与Ｐｒ均表现出明显的日变化,Ｐｎ日变化曲线里双峰型,中午前后降低。Ｐｒ随日照强度的增加而增加,至上午１１时左右达最大值,然后缓慢下降。普通空气及低氧空气中的ＡＱＹ均在中午前后降低。ＳＯＤ活性也有明显的日变化,最高值出现在下午１６时左右。强光下喷施ＳＯＤ抑制剂ＤＤＴＣ明显降低Ｐｎ及低氧空气中的ＡＱＹ;而在低于叶片光合作用饱和光强下喷施同样浓度ＤＤＴＣ则对Ｐｎ及低氧空气中的ＡＱＹ无明显影响。中午前后ＳＯＤ活性及Ｐｒ的增加对于保护光合机构免受强光的破坏具有重要意义。     

Photosynthetic acclimation to variability in the light environment of early and late successional plants

Summary Fourteen plant species from early-, mid-, and late-successional habitats were grown for a period of 25 to 50 days in each of two light environments, i.e. full sunlight and in deep shade. The rate of photosynthesis for newly formed leaves was measured as a function of light intensity for plants from each light environment. Photosynthetic flexibility, measured as the difference in response between sun- and shade-grown plants, was determined for each of 5 parameters including dark respiration, quantum yield, light compensation, half-saturating irradiance for photosynthesis, and the photosynthetic rate at 1,400 E m^-2 s^-1. We found photosynthetic flexibility to be high for early successional annuals, intermediate for midsuccessional species, and low for late successional species.     

Photosynthetic responses to dynamic light under field conditions in six tropical rainforest shrubs occuring along a light gradient

 We examined in the field the photosynthetic utilization of fluctuating light by six neotropical rainforest shrubs of the family Rubiaceae. They were growing in three different light environments: forest understory, small gaps, and clearings. Gas exchange techniques were used to analyse photosynthetic induction response, induction maintenance during low-light periods, and lightfleck (simulated sunfleck) use efficiency (LUE). Total daily photon flux density (PFD) reaching the plants during the wet season was 37 times higher in clearings than in the understory, with small gaps exhibiting intermediate values. Sunflecks were more frequent, but shorter and of lower intensity in the understory than in clearings. However, sunflecks contributed one-third of the daily PFD in the understory. Maximum rates of net photosynthesis, carboxylation capacity, electron transport, and maximum stomatal conductance were lower in understory species than in species growing in small gaps or clearings, while the reverse was true for the curvature factor of the light response of photosynthesis. No significant differences were found in the apparent quantum yield. The rise of net photosynthesis during induction after transfer from low to high light varied from a hyperbolic shape to a sigmoidal increase. Rates of photosynthetic induction exhibited a negative exponential relationship with stomatal conductance in the shade prior to the increase in PFD. Leaves of understory species showed the most rapid induction and remained induced longer once transferred to the shade than did leaves of medium- or high-light species. LUE decreased rapidly with increasing lightfleck duration and was affected by the induction state of the leaf. Fully induced leaves exhibited LUEs up to 300% for 1-s lightflecks, while LUE was below 100% for 1–80 s lightflecks in uninduced leaves. Both induced and uninduced leaves of understory species exhibited higher LUE than those of species growing in small gaps or clearings. However, most differences disappeared for lightflecks 10 s long or longer. Thus, understory species, which grew in a highly dynamic light environment, had better capacities for utilization of rapidly fluctuating light than species from habitats with higher light availability. Received: 4 January 1997 / Accepted: 28 April 1997     

小麦叶片光合作用与其渗透调节能力的关系

在缓慢干旱条件下,小麦叶片渗透调节能力在一定范围内随胁迫程度的加剧而增加,而在快速干旱下,渗透调节能力丧失。小麦叶片通过渗透调节使光合速率和气孔导度对水分胁迫的敏感性降低,叶片维持较高的电子传递能力、RuBP羧化酶活性和叶绿体光合能量转换系统活性,并推迟了小麦叶片光合速率受气孔因素限制向叶肉细胞光合活性限制转变的时间。