田间冬小麦叶片光合午休过程中的非气孔限制

1　引　　言作物的光合午休是近几年作物栽培生理研究的热点之一 ,研究大都从生态、生理的各个方面阐述光合午休的成因与机理 ,如土壤水分降低 ,ＣＯ2 浓度下降 ,ＶＰＤ过高 ,气孔限制与非气孔限制 ,羧化效率降低等[4 ,6,10 ] .其中有关作物光合午休过程中的非气孔限制问题 ,多数研究仅根据Ｆａｒｑｕｈａｒ判据[2 ] 判断发生了非气孔限制即止 ,至于非气孔限制的具体内容 ,多见于讨论 ,缺乏直接的实验证据 .本文对大田生态条件下 ,冬小麦生育后期旗叶光合午休过程中的非气孔限制问题进行了较深入的研究 .2　材料与方法2 1　供试材料选用冬…     

Stomatal closure induced by high vapor pressure deficit limited midday photosynthesis at the canopy top of <Emphasis Type="Italic">Fagus crenata</Emphasis> Blume on Naeba mountain in Japan

Diurnal changes in gas exchange, chlorophyll fluorescence and leaf water potential (_leaf) were measured to determine the environmental and physiological factors that limit carbon gain in the horizontal leaves of Fagus crenata Blume at the canopy top. Although midday depression of the net CO₂ assimilation rate (A_n) and stomatal conductance (gH₂O) were clearly evident on a fine day, the potential quantum yield of PS II (F_v/F_m) was fairly constant around 0.83 throughout the day. This result indicates that the leaves at the canopy top do not suffer from chronic photoinhibition, and the excess energy is dissipated safely. Large reversible increases in non-photochemical quenching (NPQ) were evident on fine days. Therefore, the non-radiative energy dissipation of excess light energy contributed to avoid chronic photoinhibition. The electron transfer rate (ETR) reached maximum during the midday depression, and thus there was no positive relation between ETR and A_n under high light conditions, indicating a high rate of photorespiration and the absence of non-stomatal effect during midday. The protective mechanisms such as non-radiative energy dissipation and photorespiration play an important role in preventing photoinhibitory damage, and stomatal limitation is the main factor of midday depression of A_n. To separate the effect of air to leaf vapor pressure deficit (ALVPD) and leaf temperature (T_leaf) on gas exchange, the dependencies of A_n and gH₂O on ALVPD and T_leaf were measured using detached branches under controlled conditions. A_n and gH₂O were insensitive to an increase in T_leaf. With the increase in ALVPD, A_n and gH₂O exhibited more than a 50% decrease even though water supply was optimum, suggesting the dominant role of high ALVPD in the midday depression of gH₂O. We conclude that midday depression of A_n results from the midday stomatal closure caused by high ALVPD.     

Forest fire in the central Himalaya: climate and recovery of trees

 A forest fire event is influenced by climatic conditions and is supported by accumulation of fuel on forest floor. After forest fire, photosynthetically active solar radiation was reduced due to accumulation of ash and dust particles in atmosphere. Post-fire impacts on Quercus leucotrichophora, Rhododendron arboreum and Lyonia ovalifolia in a broadleaf forest were analysed after a wild fire. Bark depth damage was greatest for L. ovalifolia and least for Q. leucotrichophora. Regeneration of saplings was observed for all the tree species through sprouting. Epicormic recovery was observed for the trees of all the species. Young trees of Q. leucotrichophora (<40 cm circumference at breast height) were susceptible to fire as evident by the lack of sprouting. Under-canopy tree species have a high potential for recovery as evident by greater length and diameter of shoots and numbers of buds and leaves per shoot than canopy species. Leaf area, leaf moisture and specific leaf area were greater in the deciduous species, with few exceptions, than in evergreen species. Received: 26 July 1995 / Revised: 1 March 1996 / Accepted: 9 September 1996     

Contributions of diffusional limitation, photoinhibition and photorespiration to midday depression of photosynthesis in Arisaema heterophyllum in natural high light

Diurnal changes in photosynthetic gas exchange and chlorophyll fluorescence were measured under full sunlight to reveal diffusional and non‐diffusional limitations to diurnal assimilation in leaves of Arisaema heterophyllum Blume plants grown either in a riparian forest understorey (shade leaves) or in an adjacent deforested open site (sun leaves). Midday depressions of assimilation rate (A) and leaf conductance of water vapour were remarkably deeper in shade leaves than in sun leaves. To evaluate the diffusional (i.e. stomatal and leaf internal) limitation to assimilation, we used an index [1–A/A₃₅₀], in which A₃₅₀ is A at a chloroplast CO₂ concentration of 350 μ mol mol ^{? 1}. A₃₅₀ was estimated from the electron transport rate (J_T), determined fluorometrically, and the specificity factor of Rubisco (S), determined by gas exchange techniques. In sun leaves under saturating light, the index obtained after the ‘peak’ of diurnal assimilation was 70% greater than that obtained before the ‘peak’, but in shade leaves, it was only 20% greater. The photochemical efficiency of photosystem II ( Δ F/F_m ′ ) and thus J_T was considerably lower in shade leaves than in sun leaves, especially after the ‘peak’. In shade leaves but not in sun leaves, A at a photosynthetically active photon flux density (PPFD) > 500 μ mol m ^{? 2} s ^{? 1} depended positively on J_T throughout the day. Electron flows used by the carboxylation and oxygenation (J_O) of RuBP were estimated from A and J_T. In sun leaves, the J_O/J_T ratio was significantly higher after the ‘peak’, but little difference was found in shade leaves. Photorespiratory CO₂ efflux in the absence of atmospheric CO₂ was about three times higher in sun leaves than in shade leaves. We attribute the midday depression of assimilation in sun leaves to the increased rate of photorespiration caused by stomatal closure, and that in shade leaves to severe photoinhibition. Thus, for sun leaves, increased capacities for photorespiration and non‐photochemical quenching are essential to avoid photoinhibitory damage and to tolerate high leaf temperatures and water stress under excess light. The increased Rubisco content in sun leaves, which has been recognized as raising photosynthetic assimilation capacity, also contributes to increase in the capacity for photorespiration.     

Temperature response of photosynthesis in transgenic rice transformed with 'sense' or 'antisense' rbcS

The responses of chlorophyll fluorescence, gas exchange rate and Rubisco activation state to temperature were examined in transgenic rice plants with 130 and 35% of the wild-type (WT) Rubisco content by transformation with rbcS cDNA in sense and antisense orientations, respectively. Although the optimal temperatures of PSII quantum efficiency and CO(2) assimilation were found to be between 25 and 32 degrees C, the maximal activation state of Rubisco was found to be between 16 and 20 degrees C in all genotypes. The Rubisco flux control coefficient was also the highest between 16 and 20 degrees C in the WT and antisense lines [>0.88 at an intercellular CO(2) pressure (Ci) of 28 Pa]. Gross photosynthesis at Ci = 28 Pa per Rubisco content in the WT between 12 and 20 degrees C was close to that of the antisense lines where high Rubisco control is present. Thus, Rubisco activity most strongly limited photosynthesis at cool temperatures. These results indicated that a selective enhancement of Rubisco content can enhance photosynthesis at cool temperatures, but in the sense line with enhanced Rubisco content Pi regeneration limitation occurred. Above 20 degrees C, the Rubisco flux control coefficient declined. This decline was associated with a decline in Rubisco activation. The activation state of Rubisco measured at each temperature decreased with increasing Rubisco content, and the slope of activation to Rubisco content was independent of temperature. We discuss the possibility that the decline in Rubisco activation at intermediate and high temperatures is part of a regulated response to a limitation in other photosynthetic processes.     

Photosynthesis of two poplar clones contrasting in O3 sensitivity

 Two clones of poplar known for their phenomenological difference in response to ozone were fumigated with 150 nl l^–1 of ozone for 5 h. In both clones the treatment significantly reduced the light-saturated rate of CO₂ uptake of recently mature leaves and this was accompanied by a decrease in stomatal conductance. Intercellular CO₂ concentration of the resistant clone increased following the fumigation. After 20 h of recovery, photosynthesis recovered completely only in the resistant clone. Electrolyte leakage of leaf disks increased in both clones to indicate damage to membranes; after the recovery time this parameter only reached values of the control in the resistant clone. The photochemical efficiency of PSII slightly decreased in the resistant clone. In the other clone, the treatment caused a decline of all chlorophyll fluorescence parameters and only some of them returned to normal values after the recovery time. The physiological response appears to be different in the two clones. In the resistant one, the most probable mechanism involved in the photosynthetic reduction was a regulatory reduction in CO₂ fixation. Also data obtained by the solute leakage indicate that in the resistant clone repair mechanisms play a role. The reduction of photosynthesis observed in the sensitive clone is related both to strong stomatal closure and to an impairment in fluorescence parameters. These alterations can indicate a general disruption at the membrane level as confirmed by the solute leakage data. Received: 30 June 1997 / Accepted: 3 September 1997     

Genetic control of macro-and micro-environmental sensitivities in Populus

Understanding the genetic mechanisms for the phenotypic plasticity and developmental instability of a quantitative trait has important implications for breeding and evolution. Two clonally replicated plantations of two 3-generation inbred pedigrees derived from the highly divergent species Populus trichocarpa and P. deltoides were used to examine the genetic control of macro- and micro-environmental sensitivities and their genetic relationships with the trait mean across two contrasting environments. For all stem-growth traits studied, the trait mean had a higher broad-sense heritability (H²) level than macroenvironmental sensitivity, both with much higher values than microenvironmental sensitivity. Genetic correlation analyses indicated that the trait mean was more or less independent of macro- or micro-environmental sensitivity in stem height. Thus, for this trait, the genetic difference in response to the two environments might be mainly due to epistasis between some regulatory loci for plasticity and loci for trait mean. However, for basal area and volume index, pleiotropic loci might be more important for their genetic differences between the two environments. No evidence was found to support Lerner’s (1954) homeostasis theory in which macro- or micro-environmental sensitivity is the inverse function of heterozygosity. Received: 8 March 1996 / Accepted: 31 May 1996     

An integrated genetic map of Populus deltoides based on amplified fragment length polymorphisms

Amplified fragment length polymorphism (AFLP) is an efficient molecular technique for generating a large number of DNA-based genetic markers in Populus. We have constructed an integrated genetic map for a Populus backcross population derived from two selected P. deltoides clones using AFLP markers. A traditional strategy for genetic mapping in outcrossing species, such as forest trees, is based on two-way pseudo-testcross configurations of the markers (testcross markers) heterozygous in one parent and null in the other. By using the markers segregating in both parents (intercross markers) as bridges, the two parent-specific genetic maps can be aligned. In this study, we detected a number of non-parental heteroduplex markers resulting from the PCR amplification of two DNA segments that have a high degree of homology to one another but differ in their nucleotide sequences. These heteroduplex markers detected have served as bridges to generate an integrated map which includes 19 major linkage groups equal to the Populus haploid chromosome number and 24 minor groups. The 19 major linkage groups cover a total of 2,927 cM, with an average spacing between two markers of 23. 3 cM. The map developed in this study provides a first step in producing a highly saturated linkage map of the Populus deltoides genome. Received: 10 September 1999 / Accepted: 3 November 1999     

Role of water relations and photosynthesis in the release of buds from apical dominance and the early reinvigoration of decapitated poplars

A decrease in xylem pressure potential starting 1 h after decapitation of young hybrid poplars ( Populus deltoides Bartr. × Populus nigra L. cv. DN22) reduced stomatal conductance and transpiration rates for the first 3 days after decapitation. This early moisture stress was alleviated 4 to 5 days after decapitation, resulting in substantial increases in stomatal aperture, transpiration and net photosynthetic rates which continued for the remainder of the one week measurement period. The results suggest the following sequence of events in the decapitated plant: After a brief moisture stress, decapitation increases moisture availability by increasing the root/shoot ratio and by reducing shoot competition for moisture. Improvement in hydration releases buds from apical dominance and increases stomatal conductance and rates of net photosynthesis. This, in turn, leads to the acceleration of growth observed when plants are reinvigorated by decapitation.     

Field comparisons of photosynthesis and leaf conductance in Ziziphus mauritiana and other fruit tree species in Zimbabwe

 Data are presented for gas exchange in exposed leaves in field grown Ziziphus mauritiana (Lamk.) at a highland site and potted seedlings at a hotter lowland site in tropical Zimbabwe, together with indigenous and locally-grown commercial fruit crops. The field trial at the highland site included local Ziziphus mauritiana, introduced Indian Z. mauritiana (cv. Umran), fig and peach. In all species assimilation was highest early in the morning, followed by a gradual decline throughout the remainder of the day. Leaf conductance followed the same trend as assimilation for fig and peach, but in Ziziphus cv. Umran and Ziziphus Musau, conductance tracked irradiance, reaching a maximum in the middle of the day. In all species, sub-stomatal CO₂ concentrations increased with declining assimilation indicating that during high irradiance, assimilation was mainly controlled by mesophyll limitations rather than conductance. At the highland site both Ziziphus cv. Umran and Musau were highly productive, with light saturated assimilation significantly higher than in either fig or peach (P < 0.01). At the warmer lowland site, assimilation and conductance in Ziziphus Musau were also higher than in other indigenous fruit tree species growing under similar conditions. Despite increased assimilation in Ziziphus, when compared to the other species, there was no increase in the assimilation ratio (ratio of assimilation/conductance) which was offset by the high conductance values. The data indicate that under conditions where water was not limiting, young Ziziphus showed no enhanced stomatal control over water loss, but was highly productive (per unit leaf area) relative to the other species. Received: 17 October 1996 / Accepted: 8 January 1997     

Drought effects on photosynthesis of the mangrove, Avicennia germinans, under contrasting salinities

 Drought effects on leaf photosynthesis of A. germinans growing under two contrasting salinities were studied in a Venezuelan fringe mangrove. During both wet and dry seasons, severe chronic-photoinhibition at predawn was not observed but strong down regulation occurred at midday during both seasons. Carbon assimilation rates (A, μmol CO₂ m⁻² s⁻¹) declined during the dry season from 11.9±1.8 to 7.0±1.5 and from 9.6±2.0 to 4.7±2.5 in plants from low and high salinity sites, respectively. Changes in carbon assimilation per unit of chlorophyll (A/Chl, mmol CO₂ mol⁻¹ Chl) were from 31.6±4.7 to 20.5±4.3 and from 21.9±4.7 to 15.2±8.2 in the low and high salinity plants, respectively. Therefore, neither changes in Chl nor seasonal differences in photoprotective down regulation could account fully for the decrease in leaf photosynthesis during drought. A reduction in CO₂ diffusion due to lowered stomatal conductance was not large enough to explain such a dramatic effect of drought on leaf photosynthesis. Stomatal response could be mitigated by the capability of A. germinans for osmotic adjustment under high salinity and/or drought. However, this intracellular salt accumulation may reduce carbon assimilation capacity further by decreasing the metabolism of leaf cells, increasing dark respiration and/or photorespiration. Received: 10 June 1998 / Accepted: 5 October 1998     

Adjustment of foliage structure and function to a canopy light gradient in two co-existing deciduous trees. Variability in leaf inclination angles in relation to petiole morphology

 Foliar inclination angles, petiole morphology and dry matter partitioning between assimilative and support biomass were studied in shade-intolerant Populus tremula L. and shade-tolerant Tilia cordata Mill. along a natural light gradient across the canopy. The leaves of sub-canopy species T. cordata were on average exposed to lower irradiances, and they were also more horizontal with greater blade inclination angles (ϕ_B, defined as the angle between the leaf fall-line and the horizon; ϕ_B was positive for the leaves inclined upwards, and negative for the leaves inclined downwards) than those in P. tremula. Seasonal average daily integrated quantum flux density (Q _int, mol m^–2 day^–1) and ϕ_B were not related in T. cordata, and only a weak negative effect of Q _int on ϕ_B was detected in P. tremula. Nevertheless, when both species were pooled, there was a strong negative relationship between Q _int and ϕ_B, implying that the leaves became progressively vertical with increasing height in the canopy. Interspecific differences in foliage inclination were mainly related to petiole morphology, in particular to petiole length, rather than to contrasting biomass investment patterns between assimilative and support tissues within the leaf. It was suggested that more horizontal leaves, resulting from the species-specific structure of petioles, partly explain the superior performance of shade-tolerant T. cordata in the understory and the sub-canopy. Received: 13 November 1997 / Accepted: 6 March 1998     

Characteristics of genetic variation in the progenies of protoplast-derived plants of rice, Oryza sativa cv Nipponbare

Genetic variation in protoplast-derived rice (Oryza sativa L.) plants was characterized using first and second generation selfed progenies. A total of 133 regenerated plants were obtained from ten protoplasts of the japonica rice cultivar Nipponbare. Sixty two regenerated plants which set enough seeds for the subsequent field tests at the next generation and were derived from five protoplasts were selected, and their selfed seeds were used as the first selfed-seed progeny generation). Fifteen plants were selected from each of the 15 lines, and their selfed seeds were used for tests at the generation. Thirty seven lines (60%) segregated plants with detrimental mutant characters of yellow-green phenotype, dwarf stature, dense and short panicle, or low seed fertility. According to the segregation patterns in the lines having mutated plants among those originated from the same protoplasts, the stages of mutation induction were estimated. Additionally, five quantitative traits were changed in almost all and lines. Varied quantitative traits of heading date, number of spikelets per panicle, and seed fertility, were in a heterozygous state. However, culm and panicle lengths showed high uniformity, whereas reduced culm and panicle lengths were caused by mutational changes in polygenes and/or multiple genes. Received: 20 March 1996 / Accepted: 21 June 1996     

Ethanol in the stems of trees

Acetaldehyde and ethanol are usually thought to be produced in plant tissues as a mechanism to tolerate hypoxic conditions. We have found acetaldehyde and ethanol to be common in the vascular cambium and in the transpiration stream of trees. In nonflooded trees, acetaldehyde and ethanol concentrations averaged 130 and 40 μ M in the cambium and 130 and 50 μ M in the xylem sap, respectively. Ethanol concentrations in the transpiration stream and the cambium increased to as much as 5 m M upon flooding. Ethanol concentrations in the vascular cambium of Populus deltoides could not be eliminated by placing logs from nonflooded trees in a pure oxygen environment for as long as 96 h, but increased by almost 3 orders of magnitude when logs were exposed to low external partial pressures of O₂. These results suggest that the vascular cambium was not hypoxic, despite the presence of acetaldehyde, ethanol and the enzymes for their synthesis.     

Assessing the impact of elevated O3 and CO2 on gas exchange characteristics of differently K supplied clonal Norway spruce trees during exposure and the following season

 Well-supplied and K-deficient 4-year-old clonal Norway spruce trees were exposed to combinations of two levels of ozone (20 and 80 nl l^{–  1} O₃) and carbon dioxide (350 and 750 μl l^{–  1} CO₂) to study the effects of possible future climate factors on gas exchange characteristics. The fumigation was performed in environmental chambers for a complete growing season. After the exposure, plants were cultivated outdoors to investigate possible recovery and delayed effects. During the exposure 1-year-old needles responded to the 80 nl l^{–  1} O₃ treatment by a sharp but transient decrease of both apparent carboxylation efficiency (CE) and maximum photosynthetic capacity (A₂₅₀₀). Elevated CO₂ also reduced CE and A₂₅₀₀. The effect became stronger in the course of the exposure and was accompanied by decreases of N and P as well as chlorophyll contents. In case of K deficiency, the acclimation response of current-year needles was even more pronounced reflecting lower sink capacities for carbon metabolites. The joint application of elevated O₃ and CO₂ resulted in the lowest values of gas exchange parameters and chlorophyll contents. At the beginning of the growing season after the exposure and under outdoor conditions, all these treatment effects disappeared in the needles which had developed during the fumigation. In the course of the development of the new flush, however, the well-supplied 1-year-old needles which had been treated with 80 nl l^{–  1} O₃ and 350 μl l^{–  1} CO₂ in the year before, exhibited a sharp decline of CE and A₂₅₀₀. Simultaneously, chlorotic mottle and bands developed. These delayed symptoms are discussed in the context of the previously published “memory” effect for O₃ (Sandermann et al. 1989). Additionally, evidence is presented that shoot development is altered in plants which had been exposed to elevated O₃. Accepted: 5 August 1996     

Effects of water and nitrogen interaction on net photosynthesis, stomatal conductance, and water-use efficiency in two hybrid poplar clones

We examined the interactions of water and nitrogen availability by subjecting two Populus clones. Tristis and Eugenei, to five soil moisture and three soil nitrogen levels. Nitrogen application significantly increased net photosynthesis and stomatal conductance of flooded Eugenei and Tristis. The onset of flooding caused partial stomatal closure. Net photosynthesis significantly declined after a longer flooding period. Emergence of adventitious roots on the submerged portions of stems in both clones seemingly helped net photosynthesis fully recover in Eugenei and partially recover in Tristis. Under the progressive drought conditions, stomatal conductance was more sensitive to drought than net photosynthesis in both clones. Addition of nitrogen to progressively drying soil induced more stomatal closure in both clones. The highest water-use efficiency was found on the high-N/severe drought zone for Eugenei, whereas it was found on the high-N/mild to moderate drought zone for Tristis.     

Is self-compatibility the main breeding system in borage (Borago officinalis L.)?

The polygenic postzygotic self-incompatible system reported in borage (Borago officinalis L.) is refuted. The behaviour of pollen tubes after self- and cross-pollination, analysis of the crossability coefficient, seed set after continuous generations of self-pollination and the effects of inbreeding depression in borage were studied. Evidence of self-compatibility was established. The influence of protandry in a self-compatible system is also discussed. Received: 15 July 1999 / Accepted: 3 November 1999     

Response of leaf surfaces and gas exchange to wind stress and acid mist in birch (Betula pubescens)

 This research demonstrates that a leaf’s response to acid mist is dependent on the integrity of the leaf cuticle and that significant differences in the structural and physiological disturbances in leaves can be attributable to different types of wind action. Betula pubescens Ehrh. plants were located at adjacent, but contrasting, sites to create different wind treatments: (i) direct wind action, (ii) indirect wind action and (iii) shelter from wind action (control). In combination with the wind treatments, acidic (pHs 5 and 3) or neutral (pH 7) mists were applied weekly. Wind action significantly increased visible leaf injury, microscopic cuticular lesions and cuticular conductance (g _c ), but reduced photosynthetic rate (P _N ) and stomatal conductance (g _s ) compared to shelter. Wind action combined with acid mist was more injurious than wind action alone, but leaves sheltered from wind action were highly resistant to the damaging effects of acid mist. Direct wind action combined with pH 3 mist resulted in the highest values of g _c and the greatest number of cuticular lesions. By contrast, indirect wind action combined with pH 3 mist induced most visible injury, but relatively low values of g _c and few microscopic cuticular lesions. Acid mist reduced P _N only when leaves had been damaged by wind action. Higher values of g _c were associated both with increases in the area of visible leaf injury and with the number of cuticular lesions. Compensatory increase in P _N of healthy tissue was evident in leaves exposed to combinations of wind action and acid mist. Received: 10 November 1997 / Accepted: 6 March 1998     

Genetic linkage mapping in Acacia mangium 1. Evaluation of restriction endonucleases, inheritance of RFLP loci and their conservation across species

Random genomic probes were used to assess levels of restriction fragment length polymorphism (RFLP) in two 2-generation outbred pedigrees of Acacia mangium Willd. Probes were evaluated for their ability to detect polymorphic loci in each pedigree and to determine the relative efficiency of different restriction enzymes in revealing polymorphisms. Sixty two percent of the probes which detected single- or low-copy number sequences revealed polymorphisms with at least one restriction enyzme. HpaII was the most efficient in detecting polymorphism among first-generation individuals. The recognition sequence of HpaII contains a CpG dimer, suggesting that cytosines in the CpG sequence may be hotspots for mutation in plant genomes, as previously reported in bacterial and mammalian genomes. Mendelian inheritance of 230 loci was demonstrated based on single-locus segregation in second-generation individuals. Less than 5% of loci showed evidence of segregation distortion. The proportion of fully informative loci (15%) was lower than previously reported in eucalypts reflecting the lower level of genetic diversity in A. mangium. The RFLP probes are suitable for the construction of a high-density genetic linkage map in A. mangium. Cross-hybridisation of the A.mangium RFLPs to DNA from species representing the three subgenera of the genus Acacia indicates that these markers could be used in breeding programs of other diploid acacias, for comparative studies of genome organisation, and for phylogenetic studies. Received: 5. June 1999 / Accepted: 30 July 1999