Elevated CO2 concentrations and stomatal density: observations from 17 plant species growing in a CO2 spring in central Italy

Stomatal density (SD) and stomatal conductance ( g _s) can be affected by an increase of atmospheric CO₂ concentration. This study was conducted on 17 species growing in a naturally enriched CO₂ spring and belonging to three plant communities. Stomatal conductance, stomatal density and stomatal index (SI) of plants from the spring, which were assumed to have been exposed for generations to elevated [CO₂], and of plants of the same species collected in a nearby control site, were compared. Stomatal conductance was significantly lower in most of the species collected in the CO₂ spring and this indicated that CO₂ effects on g _s are not of a transitory nature but persist in the long term and through plant generations. Such a decrease was, however, not associated with changes in the anatomy of leaves: SD was unaffected in the majority of species (the decrease was only significant in three out of the 17 species examined), and also SI values did not vary between the two sites with the exception of two species that showed increased SI in plants grown in the CO₂-enriched area. These results did not support the hypothesis that long-term exposure to elevated [CO₂] may cause adaptive modification in stomatal number and in their distribution.     

幼龄胡杨气孔行为对土壤质地和地下水埋深变化的响应

气孔调节是植物适应水分条件变化的关键途径,研究多变生境中植物气孔行为对认识植物的适应具有重要意义。洪水漫溢新形成的河漫滩是胡杨更新的自然生境,其土壤质地和地下水埋深具高度时空异质性。已有研究主要集中于胡杨对地下水埋深变化的生理生态响应,而对土壤质地与地下水变化交互作用影响植物水分关系的认识不足。通过设置土壤质地(砂土(S₁)、砂壤土(S₂)、黏壤土(S₃)与地下水埋深(W₁(30 cm)、W₂(60 cm)、W₃(90 cm))交互试验模拟幼龄胡杨自然生境,观测分析了不同条件下胡杨气孔导度(G_s)、气孔导度斜率(g₁)、光合的气孔限制(L_s)的变化。研究结果表明：(1)胡杨气孔行为对地下水变化的响应受土壤质地影响;(2)相同地下水埋深时不同土质间G_s具显著差异,W₁时S₂与S₃的G_s...     

Stomatal responses to CO2 during a diel Crassulacean acid metabolism cycle in Kalanchoe daigremontiana and Kalanchoe pinnata

To investigate the diurnal variation of stomatal sensitivity to CO₂, stomatal response to a 30 min pulse of low CO₂ was measured four times during a 24 h time-course in two Crassulacean acid metabolism (CAM) species Kalanchoe daigremontiana and Kalanchoe pinnata , which vary in the degree of succulence, and hence, expression and commitment to CAM. In both species, stomata opened in response to a reduction in p CO₂ in the dark and in the latter half of the light period, and thus in CAM species, chloroplast photosynthesis is not required for the stomatal response to low p CO₂. Stomata did not respond to a decreased p CO₂ in K. daigremontiana in the light when stomata were closed, even when the supply of internal CO₂ was experimentally reduced. We conclude that stomatal closure during phase III is not solely mediated by high internal p CO₂, and suggest that in CAM species the diurnal variability in the responsiveness of stomata to p CO₂ could be explained by hypothesizing the existence of a single CO₂ sensor which interacts with other signalling pathways. When not perturbed by low p CO₂, CO₂ assimilation rate and stomatal conductance were correlated both in the light and in the dark in both species.     

Stomatal sensitivity to vapour pressure difference over a subambient to elevated CO2 gradient in a C3/C4 grassland

In the present study the response of stomatal conductance (g_s) to increasing leaf‐to‐air vapour pressure difference (D) in early season C₃ (Bromus japonicus) and late season C₄ (Bothriochloa ischaemum) grasses grown in the field across a range of CO₂ (200–550 µmol mol^?1) was examined. Stomatal sensitivity to D was calculated as the slope of the response of g_s to the natural log of externally manipulated D (dg_s/dlnD). Increasing D and CO₂ significantly reduced g_s in both species. Increasing CO₂ caused a significant decrease in stomatal sensitivity to D in Br. japonicus, but not in Bo. ischaemum. The decrease in stomatal sensitivity to D at high CO₂ for Br. japonicus fit theoretical expectations of a hydraulic model of stomatal regulation, in which g_s varies to maintain constant transpiration and leaf water potential. The weaker stomatal sensitivity to D in Bo. ischaemum suggested that stomatal regulation of leaf water potential was poor in this species, or that non‐hydraulic signals influenced guard cell behaviour. Photosynthesis (A) declined with increasing D in both species, but analyses of the ratio of intercellular to atmospheric CO₂ (C_i/C_a) suggested that stomatal limitation of A occurred only in Br. japonicus. Rising CO₂ had the greatest effect on g_s and A in Br. japonicus at low D. In contrast, the strength of stomatal and photosynthetic responses to CO₂ were not affected by D in Bo. ischaemum. Carbon and water dynamics in this grassland are dominated by a seasonal transition from C₃ to C₄ photosynthesis. Interspecific variation in the response of g_s to D therefore has implications for predicting seasonal ecosystem responses to CO₂.     

鸡蛋花(夹竹桃科)花表皮气孔的初步研究

对鸡蛋花花表皮的气孔进行初步研究,结果发现:花冠裂片的上表皮没有气孔的分布;花冠裂片的下表皮则有气孔的分布。当花冠裂片长度1.5cm时,气孔密度最大,且极显著地高于花冠裂片长度为2.0、2.5、3.5cm和4.0cm时的气孔密度。长度为3.0cm的花冠裂片的气孔指数最大,与花冠裂片长度为1.0、2.0、3.5cm和4.0cm时的气孔指数的差异均达极显著水平。在花冠筒长度为0.3cm和0.4cm时,没发现气孔;当花冠筒生长到0.5cm时开始出现气孔。花冠筒长度为0.6cm时,气孔密度最大,且极显著地高于其它长度花冠筒的气孔密度。花冠筒长为0.6、1.1cm和1.3cm时的气孔指数均极显著地大于长度为0.5cm花冠筒的气孔指数。花冠裂片和花冠筒下表皮的普通表皮细胞都呈不规则的多边形,保卫细胞呈半月形。     

植物气孔对大气CO2浓度和温度升高的反应——基于在CO2浓度和温度梯度中生长的10种植物的观测

许多研究表明 ,大气 CO2 浓度 ([CO2 ])的升高会导致植物气孔密度 (Stom atal Density,SD)和气孔指数 (Stom atal Index,SI)降低。这一关系成为推测地质历史时期大气 [CO2 ]变化的重要古生物指标之一。但是 ,[CO2 ]不是唯一影响 SD和 SI的环境因素。研究利用温度梯度和温度 [CO2 ]梯度技术 ,以 7种美国中西部地区弃耕地常见草本植物和 3种美国东部落叶阔叶林优势木本植物为材料 ,其中草本包含豆科、非豆科 C3和 C4 功能型 ,就它们的 SD,SI,表皮细胞密度 (Epidermal Cell Density,ECD)和气孔孔径长度 (Stomatal Aperture L ength,APL)对 [CO2 ]和温度升高的反应进行了研究。结果表明 ,沿 [CO2 ]梯度 ,所研究物种的 SD比 SI反应敏感 ,SD显示出与 [CO2 ]正相关、负相关和无显著相关性 ,SI显示出与 [CO2 ]正相关和无显著相关性 ;沿温度梯度 ,所研究物种的 SI比 SD反应敏感 ,SI显示出与温度正相关、负相关和无显著相关性 ,SD显示出与温度正相关和无显著相关性。 ECD和 APL对 [CO2 ]和温度梯度也有不同的响应。这说明 ,除 [CO2 ]外 ,温度也对 SD,SI,ECD和 APL有显著的影响。所以在用气孔特征重建地质历史时期 [CO2 ]的变化趋势时 ,除准确建立气孔参数与 [CO2 ]关系外 ,还应考虑大气温度对这一关系的影响     

Stomatal and non-stomatal limitation to photosynthesis in two trembling aspen (Populus tremuloides Michx.) clones exposed to elevated CO2 and/or O3

Leaf gas exchange parameters and the content of ribulose‐1,5‐bisphosphate carboxylase/oxygenase (Rubisco) in the leaves of two 2‐year‐old aspen (Populus tremuloides Michx.) clones (no. 216, ozone tolerant and no. 259, ozone sensitive) were determined to estimate the relative stomatal and mesophyll limitations to photosynthesis and to determine how these limitations were altered by exposure to elevated CO₂ and/or O₃. The plants were exposed either to ambient air (control), elevated CO₂ (560 p.p.m.) elevated O₃ (55 p.p.b.) or a mixture of elevated CO₂ and O₃ in a free air CO₂ enrichment (FACE) facility located near Rhinelander, Wisconsin, USA. Light‐saturated photosynthesis and stomatal conductance were measured in all leaves of the current terminal and of two lateral branches (one from the upper and one from the lower canopy) to detect possible age‐related variation in relative stomatal limitation (leaf age is described as a function of leaf plastochron index). Photosynthesis was increased by elevated CO₂ and decreased by O₃ at both control and elevated CO₂. The relative stomatal limitation to photosynthesis (l_s) was in both clones about 10% under control and elevated O₃. Exposure to elevated CO₂ + O₃ in both clones and to elevated CO₂ in clone 259, decreased l_s even further – to about 5%. The corresponding changes in Rubisco content and the stability of C_i/C_a ratio suggest that the changes in photosynthesis in response to elevated CO₂ and O₃ were primarily triggered by altered mesophyll processes in the two aspen clones of contrasting O₃ tolerance. The changes in stomatal conductance seem to be a secondary response, maintaining stable C_i under the given treatment, that indicates close coupling between stomatal and mesophyll processes.     

Leaf stomatal responses to vapour pressure deficit under current and CO2-enriched atmosphere explained by the economics of gas exchange

Using the economics of gas exchange, early studies derived an expression of stomatal conductance ( g ) assuming that water cost per unit carbon is constant as the daily loss of water in transpiration ( f _e) is minimized for a given gain in photosynthesis ( f _c). Other studies reached identical results, yet assumed different forms for the underlying functions and defined the daily cost parameter as carbon cost per unit water. We demonstrated that the solution can be recovered when optimization is formulated at time scales commensurate with the response time of g to environmental stimuli. The optimization theory produced three emergent gas exchange responses that are consistent with observed behaviour: (1) the sensitivity of g to vapour pressure deficit ( D ) is similar to that obtained from a previous synthesis of more than 40 species showing g to scale as 1 −  m  log( D ), where m   ∈  [0.5,0.6], (2) the theory is consistent with the onset of an apparent 'feed-forward' mechanism in g , and (3) the emergent non-linear relationship between the ratio of intercellular to atmospheric [CO₂] ( c _i/ c _a) and D agrees with the results available on this response. We extended the theory to diagnosing experimental results on the sensitivity of g to D under varying c _a.     

菜心下胚轴气孔的特征及其动态变化

以"油青60天"和"四九-19"2个菜心品种为材料,研究了菜心下胚轴气孔特征、气孔密度和气孔指数在2年内随不同发育时间的变化情况.结果显示:菜心种子萌发后第3天,下胚轴表皮上已有椭圆形或狭长形的气孔分布;菜心下胚轴普通表皮细胞为长条形,排列紧密."油青60天"和"四九-19"下胚轴的平均气孔密度均先下降后上升,分别在第...     

Effect of elevated CO2 on the stomatal distribution and leaf physiology of Alnus glutinosa

Variation in stomatal development and physiology of mature leaves from Alnus glutinosa plants grown under reference (current ambient, 360 μmol mol⁻¹ CO₂) and double ambient (720 μmol mol⁻¹ CO₂) carbon dioxide (CO₂) mole fractions is assessed in terms of relative plant growth, stomatal characters (i.e. stomatal index and density) and leaf photosynthetic characters. This is the first study to consider the effects of elevated CO₂ concentration on the distribution of stomata and epidermal cells across the whole leaf and to try to ascertain the cause of intraleaf variation. In general, a doubling of the atmospheric CO₂ concentration enhanced plant growth and significantly increased stomatal index. However, there was no significant change in relative stomatal density. Under elevated CO₂ concentration there was a significant decrease in stomatal conductance and an increase in assimilation rate. However, no significant differences were found for the maximum rate of carboxylation ( V _cmax) and the light saturated rate of electron transport ( J _max) between the control and elevated CO₂ treatment.     

In vitro chromosome doubling of nineZantedeschia cultivars

Tetraploid plants have been produced from nineZantedeschia cultivars usingin vitro colchicine treatment. Rapidly-multiplying shoot cultures were treated on a medium containing 0.05% colchicine for 1, 2 or 4 days to induce chromosome doubling. Following the treatment, most shoots were killed but the surviving shoots were multiplied for several subcultures. These shoots were then rootedin vitro and transferred to a greenhouse. Plants were screened 2 months later by measuring stomatal length, and 110 out of 565 plants were selected as putative tetraploids with a stomatal length significantly greater than in diploid control plants. Chromosome counts were carried out on root tips from 44 plants and confirmed that 38 were tetraploids, 2 were chimeras (predominantly tetraploid with a few octoploid cells), and 4 were diploids. Stomatal length has been rechecked in mature tetraploid plants of the cultivar Black Magic, demonstrating that stomatal length is a good indicator of ploidy level inZantedeschia. This study has shown that multiplying colchicine-treated shootsin vitro for several subcultures prior to transfer to soil produced very few chimeras. The stomatal length measurements are non-destructive and allow the rapid screening of a population for tetraploids.     

A critical appraisal of a combined stomatal-photosynthesis model for C3 plants

Gas-exchange measurements on Eucalyptus grandis leaves and data extracted from the literature were used to test a semi-empirical model of stomatal conductance for CO₂ g_Sc=g_o+a₁A/(c_s-I) (1+D_s/D_o)] where A is the assimilation rate; D_s and c_s are the humidity deficit and the CO₂ concentration at the leaf surface, respectively; g₀ is the conductance as A → 0 when leaf irradiance → 0; and D₀ and a₁ are empirical coefficients. This model is a modified version of g_sc=a₁A h_s/c_s first proposed by Ball, Woodrow & Berry (1987, in Progress in Photosynthesis Research, Martinus Mijhoff, Publ., pp. 221–224), in which h_s is relative humidity. Inclusion of the CO₂ compensation point, τ, improved the behaviour of the model at low values of c_s, while a hyperbolic function of D_s for humidity response correctly accounted for the observed hyperbolic and linear variation of g_sc and c_i/c_s as a function of D_s, where C_i is the intercellular CO₂ concentration. In contrast, use of relative humidity as the humidity variable led to predictions of a linear decrease in g_sc and a hyperbolic variation in c_i/c_s as a function of D_s, contrary to data from E. grandis leaves. The revised model also successfully described the response of stomata to variations in A, D_s and c_s for published responses of the leaves of several other species. Coupling of the revised stomatal model with a biochemical model for photosynthesis of C₃ plants synthesizes many of the observed responses of leaves to light, humidity deficit, leaf temperature and CO₂ concentration. Best results are obtained for well-watered plants.     

田野菟丝子寄生对薇甘菊气孔行为的影响

田野菟丝子寄生已成为当前控制薇甘菊入侵蔓延的最佳生物防治手段之一。应用扫描电子显微镜技术,观察并统计分析了田野菟丝子寄生对薇甘菊叶片气孔形态参数的影响,为了解寄生对薇甘菊生理特征的影响提供一定的形态学依据。结果表明,田野菟丝子寄生对薇甘菊叶片的气孔长宽比和气孔面积产生显著影响:对照薇甘菊叶片的气孔长宽比和气孔面积表现为双峰日变化动态;而被寄生的薇甘菊的叶片气孔长宽比和气孔面积则表现为单峰日变化规律。对照薇甘菊气孔的长宽比分别在12:00和10:00,气孔面积分别在10:00和14:00达到日变化中的最大值与最小值;在15:00到16:00气孔长宽比和气孔面积再一次达到一个峰值,使气孔长宽比和气孔面积的日波动形成一个双峰曲线。被寄生的薇甘菊的叶片气孔长宽比和气孔面积则表现出单峰曲线,峰值分别出现在12:00和14:00。田野菟丝子寄生对薇甘菊的气孔密度没有产生显著影响。     

Ontogenetic changes in stomatal size and conductance of sunflowers

The ontogenetic changes in stomatal size, frequency and conductance (g_s) on abaxial and adaxial leaf surfaces of sunflower plants (Helianthus annuus L. Russian Mammoth) were examined under controlled environmental conditions. The stomatal frequency on the adaxial and abaxial leaf surfaces decreased with leaf ontogeny and insertion level. The ratio of adaxial to abaxial stomatal frequency did not change with leaf ontogeny and insertion level, and 42–44% of total stomata was apportioned to the adaxial surface. Ontogenetic changes in stomatal pore length were detected and increased with ontogenesis. The stomatal length of both leaf surfaces had linear relationships with leaf area. Ontogenetic changes in g_s were similar between the two surfaces. However the adaxial g_s was lower than abaxial g_s in leaves of higher insertion levels. Conductance had a linear relationship with width x frequency but not with pore area.     

Stomatal behaviour, photosynthesis and transpiration under rising CO2

Definitions of the variables used and the units are given in Table 1

The literature reports enormous variation between species in the extent of stomatal responses to rising CO₂. This paper attempts to provide a framework within which some of this diversity can be explained. We describe the role of stomata in the short-term response of leaf gas exchange to increases in ambient CO₂ concentration by developing the recently proposed stomatal model of Jarvis & Davies (1998 ). In this model stomatal conductance is correlated with the functioning of the photosynthetic system so that the effects of increases in CO₂ on stomata are experienced through changes in the rate of photosynthesis in a simple and mechanistically transparent way. This model also allows us to consider the effects of evaporative demand and soil moisture availability on stomatal responses to photosynthesis and therefore provides a means of considering these additional sources of variation. We emphasize that the relationship between the rate of photosynthesis and the internal CO₂ concentration and also drought will have important effects on the relative gains to be achieved under rising CO₂.  

  Table 1 . Abbreviations     

20.

Blue light inhibits stomatal development in soybean isolines containing kaempferol-3-O-2^G-glycosyl-gentiobioside (K9), a unique flavonoid glycoside     

L. Liu-Gitz  S. J. Britz  & W. P. Wergin 《Plant, cell & environment》2000,23(8):883-891

Stomata have a fundamental role in controlling plant photosynthesis and transpiration, but very little is known about factors controlling stomatal differentiation and development. Lines of soybean that contain a specific flavonol glycoside, kaempferol‐3‐O‐2‐glycosyl‐gentiobioside (K9), as well as greatly reduced stomatal density, especially on the adaxial epidermis, have been identified. The specific effects of blue light photoreceptors on stomatal development in K9 lines and their isoline pairs containing no K9 were studied. Low irradiances of blue light (7% of total photosynthetically active radiation) added to high irradiances from low‐pressure sodium lamps strongly inhibited stomatal development on the adaxial epidermis of K9 lines, but not in isoline pairs differing putatively in only one gene and lacking K9. Overall, blue light slightly increased stomatal density on the abaxial epidermis in all isolines, demonstrating differential regulation of stomatal development in the upper and lower epidermis. Blue light also caused an increase in leaf area in all isolines, indicating that changes in stomatal density were not the non‐specific result of alterations in leaf area. Morphological studies revealed that the blue light‐induced reduction in stomatal density in K9 lines was due to reduced stomatal initiation as well as aborted or abnormal stomatal development. As the phytochrome photostationary state was kept constant, the results indicate that one or more blue light receptors are involved in the control of stomatal development. This system should be useful for the study of mechanisms controlling stomatal development, even if the photo‐inhibitory response is unique to K9 lines.     

Blue light inhibits stomatal development in soybean isolines containing kaempferol-3-O-2G-glycosyl-gentiobioside (K9), a unique flavonoid glycoside

Stomata have a fundamental role in controlling plant photosynthesis and transpiration, but very little is known about factors controlling stomatal differentiation and development. Lines of soybean that contain a specific flavonol glycoside, kaempferol‐3‐O‐2‐glycosyl‐gentiobioside (K9), as well as greatly reduced stomatal density, especially on the adaxial epidermis, have been identified. The specific effects of blue light photoreceptors on stomatal development in K9 lines and their isoline pairs containing no K9 were studied. Low irradiances of blue light (7% of total photosynthetically active radiation) added to high irradiances from low‐pressure sodium lamps strongly inhibited stomatal development on the adaxial epidermis of K9 lines, but not in isoline pairs differing putatively in only one gene and lacking K9. Overall, blue light slightly increased stomatal density on the abaxial epidermis in all isolines, demonstrating differential regulation of stomatal development in the upper and lower epidermis. Blue light also caused an increase in leaf area in all isolines, indicating that changes in stomatal density were not the non‐specific result of alterations in leaf area. Morphological studies revealed that the blue light‐induced reduction in stomatal density in K9 lines was due to reduced stomatal initiation as well as aborted or abnormal stomatal development. As the phytochrome photostationary state was kept constant, the results indicate that one or more blue light receptors are involved in the control of stomatal development. This system should be useful for the study of mechanisms controlling stomatal development, even if the photo‐inhibitory response is unique to K9 lines.