边界层阻力在叶片气体交换过程中的作用

作者对现有的气体交换法的叶室作了改进。从实验上检验了作者根据电学类比分析和模拟检验得到的结论：在已有的气体交换测定装置中,空气流速不变,叶片的边界层导度也固定不变,所得到的结论用于边界层导度变化的情形时,会造成比较大的误差。现加一障碍物使得流过叶片的风速减小,从而改变叶片的边界层导度。通过测量在不同边界层导度下同一叶片的光强曲线可以考察不同边界层导度下的气孔导度、光合速率、蒸腾速率以及它们之间的相互关系。实验结果与作者以前的电学类比分析和模拟得到的结论是基本一致的     

The Electrical Analogy Analysis and Simulation Testing of the Relationship Between Net Photosynthetic Rate and Stomatal Conductance

The hypothesis about the linear relationship between net photosynthetic rate and stomatal conductance, which had formed the basis of the empirical model proposed by Ball et al, was analyzed by analogy with electrical resistance and simulation testing. The results showed that the relationship between stomatal conductance and net photosynthetic rate was non-linear, but approaches a linear one when there was a large the boundary layer conductance of leaf. In addition, the way in which the ratio of CO2 concentrations in and out of the stomata changes with light and the influence of boundary layer conductance on the ratio had also been simulated.     

Simulation of the Physiological Responses of C3 Plant Leaves to Environmental Factors by a Model Which Combines Stomatal Conductance,Photosynthesis and Transpiration

Transpiration element is included in the integrated stomatal conductance-photosynthesis model by considering gaseous transfer processes, so the present model is capable to simulate the influence of boundary layer conductance. Leuning in his revised Ball' s model replaced relative humidity with VPDs(the vapor pressure deficit from stomatal pore to leaf surface) and thereby made the relation with transpiration more straightforward, and made it possible for the regulation of transpiration and the influence of boundary layer conductance to be integrated into the combined model. If the differences in water vapor and CO2 concentration between leaf and ambient air are considered, VPDs, the evaporative demand, is influenced by stomatal and boundary layer conductance. The physiological responses of photosynthesis, transpiration, and stomatal function, and the changes of intercellular CO2 and water use efficiency to environmental factors, such as wind speed, photon flux density, leaf temperature and ambient CO2, are analyzed. It is shown that ff the boundary layer conductance drops to a level comparable with stomatal conductance, the results of simulation by the model presented here differ significantly from those by the previous model, and, in some cases, are more realistic than the latter.     

净光合速率与气孔导度相互关系的电学类比分析和模拟研究

对Ｂａｌｌ等的气孔模型所依据的实验事实之一（净光合速率与气孔导度呈线性关系）作了只考虑各阻力与总通量之间关系的电学类比分析和模拟检验。从电学类比分析得到的关系式可以看出,净光合速率与气孔导度的关系是非线性的,只有当叶片的边界层导度比较大时,两者的关系才接近线性。模拟得到的结论也是一样的。另外还模拟了气孔内外ＣＯ２ 浓度之比随光强的变化以及边界层导度的影响,模拟结果可以解释已有的实验结果     

红树林植物叶片形态和解剖特征对叶肉导度、叶片导水率的影响

叶肉导度和叶片导水率是影响光合作用的两个重要过程,叶肉导度通过影响从气孔下腔到Rubisco酶位点的二氧化碳浓度梯度直接影响光合作用,而叶片导水率则通过影响水分供应或气孔行为来影响光合作用,然而对这两个生理过程之间的协同性研究较少。本研究选择9种红树林植物为研究对象,探讨盐生环境下植物叶肉导度和叶片导水率的协同性及其与叶片解剖结构特征之间的相关性。结果表明,9种红树林植物叶片导水率(0.78~5.83 mmol·m~(-2)·s~(-1)·MPa-1)、叶肉导度(0.06~0.36 mol·m~(-2)·s~(-1))、最大光合速率(7.23~23.71μmol·m~(-2)·s~(-1))等特征的差别较大;叶肉导度与最大光合速率呈显著正相关,而与比叶重无显著相关性,其原因是由于比叶重与叶片厚度、叶片密度不存在相关性;叶脉密度与气孔密度呈较强的相关性,说明红树林植物叶片水分运输与散失相关的叶片结构之间存在协同关系;叶片导水率不受叶脉密度影响,并且与叶肉导度、最大光合速率也不存在相关性,这很可能与红树林植物叶片的肉质化、有发达的储水组织有关,体现了红树林植物叶片结构和功能的特殊性。     

光合作用-蒸腾作用-气孔导度的耦合模型及C_3植物叶片对环境因子的生理响应(英文)

以叶片的气体传输过程为基础,将蒸腾作用包括在以往光合作用气孔导度的耦合模型中,建立了光合作用蒸腾作用气孔导度的耦合模型。该模型可以模拟边界层导度对生理过程的影响。模拟了Ｃ３植物叶片对环境因子,如光照、温度、湿度、边界层导度和ＣＯ２浓度等的生理响应（光合作用、蒸腾作用、气孔导度）以及Ｃｉ和水分利用效率的变化。在环境因子变化于较大范围的情况下,模拟结果符合许多实验结论。     

Genotypic variation for drought tolerance in cotton (Gossypium hirsutum L.): Leaf gas exchange and productivity

Although water-limited environments are detrimental to cotton growth and productivity worldwide, identification of cotton (Gossypium hirsutum L.) genotypes that are less sensitive to drought may improve productivity in drought prone areas. The objective of the study was to assess genotypic variation for drought tolerance in cotton varieties using physiological attributes as selection criteria, and to determine the relationship of physiological attributes with productivity traits. The association of target physiological traits for drought tolerance (photosynthetic rate (P_n), stomatal conductance (g_s), and transpiration rate (E)) with productivity traits under well-watered (W₁) and water-limited (W₂) regimes was analyzed using 32 public cotton cultivars/bred lines in two field experiments conducted during the normal cotton growing seasons 2003 and 2004. Seed cotton yield (SCY) and biological yield (BY) were markedly affected under W₂ regime in all cultivars except the outstanding performance of CIM-1100 and RH-510 proving their superiority to other cultivars in drought tolerance. Conversely, FH-901, FH-634, and FH-2000 were high yielding under W₁ regime; however, exhibited a sharp decline in yield under W₂ regime. A positive correlation between SCY and BY under water stress (r=0.44 in 2003; r=0.69 in 2004) indicates that BY is also a primary determinant of SCY under water stress and genetic improvement of BY under water-limited environment may also improve SCY. P_n, g_s, and E were significantly reduced by water stress. Substantial genotypic variation for gas exchange attributes existed among the cotton cultivars. A positive association (P<0.01) was observed between g_s and E under both regimes in both years indicating the prevalence of stomatal control of transpiration. The positive association (P<0.01) between P_n and g_s in both years in W₂ regime indicates also a major role of stomatal effects in regulating leaf photosynthesis under water-limited conditions. P_n was significantly correlated with SCY (P<0.01) and BY (P<0.05 in 2003; P<0.01 in 2004) in W₂ regime; however, the level of these associations was not significant in W₁ regime. These findings demonstrate that association of P_n with productivity is effective under water-limited environment and may be useful as a selection criterion in breeding programs with the objective of improving drought tolerance and SCY under water-limited environments. Moreover, association between SCY and BY under water stress suggests that genetic improvement of BY under water stress may also improve SCY.     

拟单性木兰不同季节断根和剪枝后蒸腾和光合特性的变化

在2005年春季、夏季和秋季,对10a生拟单性木兰(Parakmeriaomeiensis)进行断根和剪枝处理,模拟移栽,用LICOR-6400测定了植株叶片的蒸腾速率、光合速率等生理指标,探讨这些生理指标对不同季节移栽成活率的影响。结果表明,春季处理后叶片能迅速关闭部分气孔,蒸腾速率和光合速率都减小,两者最低时约为对照的60%,生理机能的减弱有利于维持地上、地下部分的平衡,因而最有利于植株成活。夏季处理后叶片气孔导度显著增加,光合速率和蒸腾速率也随之增大,最高时约为对照的2倍左右,此时,若主要依靠剪枝来降低水分消耗,很难保证水分代谢平衡,移栽成活率低。秋季处理后植株叶片气孔导度高于对照,光合速率和蒸腾速率也增加,但增加幅度相对夏季较小,前期最高约增加40%,此时移栽可以通过适当的剪枝和增加土壤水分含量等措施提高成活率。可见,与水分相关的生理机能的调节机制,决定了移栽后的成活水平,即处理后能迅速调节自身生理机能、减少水分消耗的植株,成活率就高,反之就低。     

Variation of Leaf Stomatal Conductance in Winter Wheat Canopy

Leaf stomatal conductance measured and analysed in the canopies of two winter wheat varieties in the field revealed that the probability of adaxial to abaxial conductance ratio followed an approximately normal distribution with a peake value of about 1.5. The ratio changed with the developmental stages being maximium at the heading stage. Leaf stomata in wheat of the upper part of the canopy were more active and showed more pronounced diurnal change of conductance than those of the lower part. Stomatal conductance decreased from top to bottom in canopy as a negative exponential function. By comparing adaxial and abaxial conductances in the apical, middle and basal parts of a leaf, the distribution of the stomatal conductances of a wheat leaf was as follows: a steady decrease from the basal part of adaxial, through the middle and apical parts of the adaxial surface turning to the apical part of abaxial, and then the middle and lastly, the basal part of abaxial. Based on values of the correlation coefficients among the various stomatal conductance and average stomatal conductance, the authors suggested that optimal apical measurement of stomatal conductance would be at the middle and apical parts and that of abaxial would be at middle and basal parts.     

土壤水分状况对花生和早稻叶片气体交换的影响

通过田间测坑试验研究了长期处于不同土壤水分状况下花生和早稻叶片气体交换的一些特点.结果表明,花生分枝期轻度和中度水分胁迫使气孔导度(Gs)和蒸腾速率(Tr)略有下降,净光合速率(Pn)和叶片水分利用效率(WUE)减小,轻度水分胁迫Gs/Tr略有上升而中度胁迫Gs/Tr变小.花生结荚期轻度和中度水分胁迫都使Gs、Tr、Gs/Tr和Pn显著降低,WUE大幅度上升.花生结荚期明显受土壤水分胁迫影响.早稻灌浆期轻度和中度水分胁迫Gs、Tr和Gs/Tr变化不显著,Pn和WUE增加,并且轻度水分胁迫下籽粒产量增加.Gs和Gs/Tr变化情况相结合可以作为作物水分胁迫程度的一个参考指标,即如果Gs和Gs/Tr同时下降则作物已经受到水分胁迫影响.     

Evaluation of physiological traits for improving drought tolerance in faba bean (<Emphasis Type="Italic">Vicia faba</Emphasis> L.)

Among grain legumes, faba bean is becoming increasingly popular in European agriculture due to recent economic and environmental interests. Faba bean can be a highly productive crop, but it is sensitive to drought stress and yields can vary considerably from season to season. Understanding the physiological basis of drought tolerance would indicate traits that can be used as indirect selection criteria for the development of cultivars adapted to drought conditions. To assess genotypic variation in physiological traits associated with drought tolerance in faba bean and to determine relationships among these attributes, two pot experiments were established in a growth chamber using genetic materials that had previously been screened for drought response in the field. Nine inbred lines of diverse genetic backgrounds were tested under adequate water supply and limited water conditions. The genotypes showed substantial variation in shoot dry matter, water use, stomatal conductance, leaf temperature, transpiration efficiency, carbon isotope discrimination (Δ¹³C), relative water content (RWC) and osmotic potential, determined at pre-flowering vegetative stage. Moisture deficits decreased water usage and consequently shoot dry matter production. RWC, osmotic potential, stomatal conductance and Δ¹³C were lower, whereas leaf temperature and transpiration efficiency were higher in stressed plants, probably due to restricted transpirational cooling induced by stomatal closure. Furthermore, differences in stomatal conductance, leaf temperature, Δ¹³C and transpiration efficiency characterized genotypes that were physiologically more adapted to water deficit conditions. Correlation analysis also showed relatively strong relationships among these variables under well watered conditions. The drought tolerant genotypes, ILB-938/2 and Melodie showed lower stomatal conductance associated with warmer leaves, whereas higher stomatal conductance and cooler leaves were observed in sensitive lines (332/2/91/015/1 and Aurora/1). The lower value of Δ¹³C coupled with higher transpiration efficiency in ILB-938/2, relative to sensitive lines (Aurora/1 and Condor/3), is indeed a desirable characteristic for water-limited environments. Finally, the results showed that stomatal conductance, leaf temperature and Δ¹³C are promising physiological indicators for drought tolerance in faba bean. These variables could be measured in pot-grown plants at adequate water supply and may serve as indirect selection criteria to pre-screen genotypes.     

荷木人工林蒸腾与冠层气孔导度特征及对环境因子的响应

为了解荷木人工林水分利用特征及与环境因子的关系,利用Granier热消散探针于2007年12月(干季)和2008年9月(湿季)对荷木(Schima superba)人工林的15株样树树干液流进行测定并结合环境因子,计算林段总蒸腾(Jd)和冠层气孔导度(gs)。结果表明:边材面积(As)越大的径级对林段总蒸腾量的贡献越大。Jd干湿季差异显著,9月和12月平均分别为21.1 g H2O s-1和7.03 g H2O s-1,显示了明显的季节变化。环境因子与Jd呈极显著相关性,相关系数依次为:光合有效辐射(PAR)水汽压亏缺(VPD)空气相对湿度气温。gs在9月和12月的最大值分别为30.8 mmol m-2s-1和19.7 mmol m-2s-1;gs与PAR呈线性正相关;当PAR1000μmol m-2s-1且VPD2kPa时,gs与VPD呈线性负相关;偏相关分析表明土壤含水量与gs没有显著的相关性,显示所研究时间内土壤水分状况对荷木人工林蒸腾没有显著影响。     

The Simulation of Leaf Conductance Responses to Changes in Environmental Water Status

A simulation model of stomatal response to change of environmental water status was set up based on the works on the mechanism of stomatal movement. The variations of leaf conductance, water potential and turgot pressure in guard cells, subsidiary cells and the other cells or tissues in leaf with leaf-air vapour pressure difference and soil water potential have been calculated by our model. The calculated results fit very well with the data from experiments. The different patterns of leaf transpiration variation with the difference between leaf-air and vapour pressure can be explained quantitatively.     

鹤山人工马占相思林水分生态研究

 通过对马占相思叶片的蒸腾速率、气孔导度及其相应环境因子的测定,探讨鹤山丘陵人工马占相思林的水分生理生态特征。结果表明：蒸腾速率和气孔导度具有明显的日变化,两者的变化趋势相似,在夏季的日变化曲线呈双峰型,冬季呈单峰型;夏季的蒸腾速率最高,在冬季最低,仅占夏季的27.6％;马占相思林的蒸腾耗水量在旱季和雨季有明显的差异,旱季的蒸腾耗水量占雨季的25.5％,年蒸腾量为1625.1mm,占同期降水量的78.3％;马占相思林的蒸腾量接近热带雨林。     

Leaf gas exchange of trees in old-growth and young secondary forest stands in Sulawesi, Indonesia

In the tropics, old-growth forests are converted to other land cover types at a high rate and young secondary forest may gain in importance. Information on associated changes in leaf gas exchange and other leaf traits can be valuable for modelling biogeochemical fluxes under altered land-use patterns. We studied in situ photosynthetic parameters and stomatal conductance for water vapour in eight abundant tree species of young secondary forest and eight tree species of natural old-growth forest in Central Sulawesi, Indonesia. In sun leaves, the average maximal stomatal conductance (g _smax) in the secondary forest (SF) species was 2.1 times higher than in the old-growth forest (OGF) species. Species with a high g _smax reduced g _s sharply when vapour pressure deficit of the air increased, whereas species with a low g _smax were much less sensitive to air humidity. For area-based photosynthetic capacity (A _max-area), the SF species had a 2.3 times higher average than the OGF species. For both, g _smax and A _max-area the variation among species was higher in the OGF than in the SF. When all tree species (n=16) are considered, species means of specific leaf area (SLA), leaf N concentration and leaf P concentration were significantly correlated with g _smax and A _max-area. The strong correlation between A _max-area and foliar P (r ²=0.8) is remarkable as the alluvial soils in the study region are rich in nutrients. If the eight OGF species are analysed separately, the only significant correlation was observed between SLA and mass-based A _max; in the SF species strong correlations were found between leaf size and A _max-area and g _smax. These results show that the conversion of old-growth forest to young secondary forest in Sulawesi significantly alters tree leaf gas exchange characteristics and that chemical and structural leaf traits can be used for the prediction of these changes. The best correlations between leaf gas exchange parameters and leaf traits were obtained by different traits in the SF species, the OGF species and the entire pool of studied species.     

茶树蒸腾特性的研究

茶树蒸腾速率和气孔导度因品种、叶位不同而异．抗旱性强的品种具有低的气孔导度;鱼叶的蒸腾速率和气孔导度接近或高于真叶．茶树叶片蒸腾速率夏、秋季最高,春季次之,冬季最低．在晴天,茶树蒸腾速率和气孔导度的日变化呈单峰型,以中午前后最高;其日变化与光量子通量密度、叶温、蒸汽压亏缺等因子显著相关;光量子通量密度对蒸腾速率和气孔导度影响较明显．     

Leaf photosynthetic traits scale with hydraulic conductivity and wood density in Panamanian forest canopy trees

We investigated how water transport capacity, wood density and wood anatomy were related to leaf photosynthetic traits in two lowland forests in Panama. Leaf-specific hydraulic conductivity (k_L) of upper branches was positively correlated with maximum rates of net CO₂ assimilation per unit leaf area (A_area) and stomatal conductance (g_s) across 20 species of canopy trees. Maximum k_L showed stronger correlation with A_area than initial k_L suggesting that allocation to photosynthetic potential is proportional to maximum water transport capacity. Terminal branch k_L was negatively correlated with A_area/g_s and positively correlated with photosynthesis per unit N, indicating a trade-off of efficient use of water against efficient use of N in photosynthesis as water transport efficiency varied. Specific hydraulic conductivity calculated from xylem anatomical characteristics (k_theoretical) was positively related to A_area and k_L, consistent with relationships among physiological measurements. Branch wood density was negatively correlated with wood water storage at saturation, k_L, A_area, net CO₂ assimilation per unit leaf mass (A_mass), and minimum leaf water potential measured on covered leaves, suggesting that wood density constrains physiological function to specific operating ranges. Kinetic and static indices of branch water transport capacity thus exhibit considerable co-ordination with allocation to potential carbon gain. Our results indicate that understanding tree hydraulic architecture provides added insights to comparisons of leaf level measurements among species, and links photosynthetic allocation patterns with branch hydraulic processes.     

模拟酸雨对柚木幼苗生长、光合与水分利用的影响

模拟pH6.5(对照)、4.5和2.5三个酸雨梯度,研究其对1a生组培柚木(TectonagrandisL.f.)幼苗生长、光合与水分利用的影响。结果表明,尽管不同处理间的各项生理指标差异不明显,但模拟酸雨对柚木形态构件参数造成较严重的影响。pH4.5和pH2.5处理组柚木基径(D)和树高(H)增长明显下降,使得D2H下降更加显著;不同处理下柚木叶片净光合速率(Pn)和蒸腾速率(E)日变化趋于一致,气孔导度(gs)日变化与对应的叶片净光合速率日变化十分相似,同时,对照与两个处理的Pn与gs之间都表现正相关(p<0.01),且在pH4.5处理表现更为显著,但是对照和两个处理的E与gs的线性关系不显著;pH4.5和2.5处理的水分利用效率(WUE)日变化趋于一致;对照胞间CO2浓度与大气CO2浓度比(Ci/Ca)均值最低,表明对照柚木对CO2利用最有效。     

淹水时水翁幼苗光合特性与不定根的关系(简报)

淹水引起水翁幼苗净光合速率、气孔导度和蒸腾速率发生变化。淹水５ｄ后,水翁幼苗净光合速率、气孔导度和蒸腾速率慢慢下降。但从淹水第３５天开始,部分水翁幼苗在淹水的茎部产生不定根,有不定根的水翁幼苗的净光合速率、气孔导度和蒸腾速率逐渐提高,到第８０天后维持在较高水平。有不定根的水翁幼苗的净光合速率、气孔导度和蒸腾速率的日进程呈双峰型,回归分析表明,净光合速率与气孔导度呈正相关（ｒ＝０．６９,Ｐ〈０．０５     

Changes in Stomatal Conductance,Transpiration and Water Use Efficiency of Ten Species Experienced in High CO2 Concentrations in Biosphere 2

The stomotal conductance, transpiration and water use efficiency (WUE) were measured using a LI-6400 portable photosynthesis system for 5 tropical rain forest species and 5 desert species in Biosphere 2, USA. All the species have experienced in very high CO2 ( > 2 200 μmol• mol- 1 ) for more than 4.5 years. The results showed that the stomatal conductance and transpiration of rain forest species decreased from ( 127.4 ± 65.6) and (2.04 ± 0.61 ) mmol• m- 2•s- 1 to (61.3 + 30.5) and ( 1.54 ± 0.65 ) mmol• m-2• s -1 respectively, while WUE increased from (2.90 ± 0.55) to (8.45 ± 2.71) μmol CO2 •mmo1-1 H2O, with CO2 increasing from 350 – 400 to 700 – 820 μmol• mol-l. For the desert species, stomatal conductance and transpiration decreased from respectively (142.8±94.6) and (2.09±0.71) mmol•m-2•s-1 to (57.7±35.8) and (1.36±0.52) mmolm-2•s-l, but WUE increased from (4.69 ± 1.39) to (9.68 ± 1.61) μmol CO2•mmo1-1 H2O, with the CO2 increase from 320 - 400 to 820 – 850 μtmol• mol- 1. The stomatal conductance, transpiration and WUE were less influenced by light intensity under high CO2 than low CO2 concentrations. Most rain forest species reached their light saturation points at light intensity of 500 μmol• m-2•s-1, while desert species at 1 000 μmol•m-2•s-1. Among different species, the desert C3 tree, Nicotiana glauca Grah., had the highest decrease in stomatal conductance and transpiration and the highest increase in WUE, by 78%, 69% and 310% respectively. The enhancement of increasing CO2 to the stomatal, transpiration and WUE of species with different photosynthesis pathway and life forms in Biosphere 2 could be concluded as: C3 species > C4 species, and desert C3 species > rain forest C3 species.